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G "^ \ 

( 1 

BULLETIN OF 

THE BRITISH MUSEUM 
(NATURAL HISTORY) 



ZOOLOGY 

Vol. 21 

19701972 



BRITISH MUSEUM (NATURAL HISTORY) 
LONDON: 1975 



DATES OF PUBLICATION OF THE PARTS 

No. I . . . -.22 December 1970 

No. 2 . . . . .28 April 1971 

No. 3 . . . . .10 September 1971 

No. 4 . . . .10 September 1971 

No. 5 . . . .21 September 1971 

No. 6 . . . . .3 December 1971 

No. 7 . . . .3 December 1971 

No. 8 ..... 4 February 1972 



Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol BS4 5NU 



CONTENTS 

ZOOLOGY VOLUME 21 




No. i. Decapod crustacean larvae collected during the International 
Indian Ocean Expedition. Families Raninidae and Homolidae. 
By A. L. RICE 

No. 2. Bats from Ethiopia collected by the Great Abbai Expedition, 
1968. By J. E. HILL and P. MORRIS (Pis. 1-3) 

No. 3. On some species of parasitic worms in the 'Discovery' collections 
obtained in the years 1925-1936. By S. MARKOWSKI 

No. 4. Mites of the genus Hypoaspis Canestrini, 1884 5. sir. and related 
forms (Acari : Mesostigmata) associated with beetles. By 
M. COSTA .......... 

No. 5. The polychaete fauna of the Solomon Islands. By P. E. GIBBS 

No. 6. Lizards and snakes from Transjordan, recently acquired by the 
British Museum (Natural History). By Y. L. WERNER (Pis. 1-6) 

No. 7. Conchoecia from the North Atlantic, the 'procera' group. By 
M. V. ANGEL 



Miscellanea .......... 

Conchoecia pseudoparthenoda (nov. sp.), a new halocyprid ostra- 

cod for the Tropical North Atlantic. By M. V. ANGEL . 

The type specimens and identity of the species described in the 

genus Lithobius by George Newport in 1844, 1845 and 1849 

(Chilopoda, Lithobiomorpha). By E. H. EASON . 

A re-examination of the chestnut-shouldered wren complex of 

Australia. By C. J. O. HARRISON ..... 

A new species of Lironeca (Isopoda ; 

on cichlid fishes in Lake Tanganyika. 

Eunice manihine sp. nov. (Polychaeta 

of the flavus-bidentate group from 

Indian Ocean. By M. R. LONGBOTTOM .... 

Original description of the gangetic dolphin, Platanista gange- 
tica, attributed to William Roxburgh. By G. PILLERI . 
A new genus and species of Sudan leech formerly confused with 
Limnatis nilotica (Hirudinidae s.l. : Hirudinea). By L. R. 
RICHARDSON ......... 

The type-species of the genera Phoxinellus, Pseudophoxinus and 
Paraphoxinus (Pisces, Cyprinidae). By E. TREWAVAS . 



Cymothoidae) parasitic 
By R. J. LINCOLN 
: Eunicidae), a member 
the western equatorial 



25 



Index to Volume 21 



67 
99 

213 

257 

285 
289 

297 
313 
329 

339 
345 

349 
359 
363 



DECAPOD CRUSTACEAN LARVAE 

COLLECTED DURING THE 
INTERNATIONAL INDIAN OCEAN 
EXPEDITION. FAMILIES RANINIDAE 
AND HOMOLID 




A. L. RICE 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 21 No. i 

LONDON : 1970 



DECAPOD CRUSTACEAN LARVAE COLLECTED 

DURING THE INTERNATIONAL INDIAN 
OCEAN EXPEDITION. FAMILIES RANINIDAE 

AND HOMOLIDAE 




BY 

ANTHONY LEONARD RICE 



Pp. 1-24; 9 Text-figures 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 21 No. i 

LONDON: 1970 



THE BULLETIN OF THE BRITISH MUSEUM 

(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 21 No. i of the Zoological 
series. The abbreviated titles of periodicals cited 
follow those of the World List of Scientific Periodicals. 



World List abbreviation 
Bull. Br. Mus. nat. Hist. (Zool.). 



Trustees of the British Museum (Natural History), 1970 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 22 December, 1970 Price 



DECAPOD CRUSTACEAN LARVAE COLLECTED 

DURING THE INTERNATIONAL INDIAN 

OCEAN EXPEDITION. FAMILIES RANINIDAE 

AND HOMOLIDAE 

By A. L. RICE 

SYNOPSIS 

Five crustacean larvae belonging to the family Raninidae and two belonging to the family 
Homolidae are described from material collected during the International Indian Ocean exped- 
dition. Probable identities of the larvae are suggested where possible. Larval evidence for 
the relationship between the Homolidae, the Raninidae and the higher Brachyura is discussed. 

INTRODUCTION 

THE larval stages of decapod crustaceans of the Indian Ocean are rather poorly 
known, few having been hatched from the adults and even fewer reared through all 
the larval stages. Consequently, only a very small proportion of larvae taken in 
the plankton in this region can be identified to species with any certainty, and in 
many cases identification even to family is difficult. 

In those decapodan families in which the larval characteristics are already well 
known an account of plankton caught material of unknown specific or even generic 
identity is of doubtful value. On the other hand, the larvae of some families are so 
poorly known that any information on the developmental stages is of value, even if 
the material on which this information is based is at the moment unidentifiable. 
Such is the case with the crab-like families Raninidae and Homolidae, and although 
the I.I.O.E. collections contain very little material of either group a report on it is 
warranted. 

Family RANINIDAE 

Larval stages belonging to some ten species of raninids have been described 
previously, but in only three cases have the larvae been definitely identified with a 
known adult. Ranina ranina (L.) larvae were hatched from the egg by Aikawa 
(1941) and the first stage described. Lyreidus tridentatus de Haan larvae were 
reared from the egg to the moult from the 5th to the 6th (last) zoeal stage by William- 
son (1965) who also had plankton caught megalopae which moulted in the laboratory 
to the first young crab stage. Finally, Knight (1968) reared larvae of Raninoides 
benedidi Rathbun taken from the plankton off the Pacific coast of Mexico, some 
specimens collected as first zoeae surviving into the early crab stages. 

Two of the above species, R. ranina and L. tridentatus, have been recorded as 
adults from the Indian Ocean, but the larvae of the other eight recorded Indian 



A. L. RICE 



TABLE i 
Indian Ocean records of adult raninid crabs 



Species 

Cosmonotus grayi 
Adams and White 



Lyreidus channeri 
Wood-Mason 



Lyreidus tridentatus 
de Haan 

Notopus dorsipes 
(Fabr.) 



Notosceles chimmonis 

Bourne 

Notosceles viaderi Ward 
Ranina ranina (L.) 



Raninoides hendersoni 

Chopra 
Raninoides personatus 

\Vhite 

Raninoides serratifrons 
Henderson 



Locality 

Persian Gulf 
Dar-es-Salaam 
Holothuria Bank, 

i335'S: i26E 
Bay of Bengal, 

21 6'3o"N: 89 2o'E 
Bay of Bengal, 9 i4'io"N: 75 46'E 
Bay of Bengal 
Andaman Sea 
"Both sides of Ceylon" 
Malabar Coast 

Dar-es-Sallaam 

N.E. Laurence Marques, Mozambique, 

2532'S: 33 2 4 'E 
Malabar Coast 
Andamans 
Zanzibar 
Mauritius 
Amirante Islands 
Seychelle Islands 
Mauritius 
Durban 
Delagoa Bay 
Zululand Coast 
Mozambique Channel, 

I 9 5'S: 36 2i'E 
Mozambique 
Mauritius 
Reunion 
Andaman Sea, 

n49'5o"N:92 52'E 
Bay of Bengal, 

off the mouth of the river Hughli 
Bay of Bengal 
Holothuria Bank 
Off Cape Negrais, Burma 

i525'N: 9 3 45'E 
Off Travancore coast, 

955'N: 7545'E 
Off southern Ceylon, 

6 2'3"N: 81 29'E 
Ceylon 
Ceylon 

Malabar Coast 
Port Shepstone, Natal 



Source 



Alcock, 1896 
Doflein, 1904 

British Museum (Nat. Hist. 

Wood-Mason, 1886 
Alcock, 1899 
Alcock, 1896 
Alcock, 1896 
Alcock, 1896 
Alcock, 1896 

Doflein, 1904 

U.S. Nat. Mus. 
Alcock, 1896 
Alcock, 1896 
Nobili, 1905 
Studer, 1882 
U.S. Nat. Mus. 
U.S. Nat. Mus. 
Ward, 1942 
Barnard, 1950 
Barnard, 1950 
Barnard, 1950 

U.S. Nat. Mus. 
Bianconi, 1851 
Bouvier, 1915 

Hoffman, 1874 

Chopra, i933b 

Chopra, I933a 
Alcock, 1896 
Henderson, 1893 

Zool. Survey India 
Zool. Survey India 

Zool. Survey India 
Laurie, 1906 
Alcock, 1896 
Alcock, 1896 
Barnard, 1950 



INDIAN OCEAN RANINID AND HOMOLID LARVAE 



D 




FIG. i. Ranina ranina (L.), stage I zoea. A, Posterior view; B, lateral view; C, frontal view; 
D, dorsal view of abdomen; E, enlarged frontal view of lateral carapace spine of left-hand 
side; F, enlarged dorsal view of postero-lateral part of telson. Bar scale represents 2.0 mm 
for A, i-o mm for B, C and D, and 0-2 mm for E and F. 



A. L. RICE 




FIG. 2. Ranina ranina (L.), stage I zoea. A, Antennule; B, antenna; C, mandibles; D, 
maxillule; E, maxilla; F, first maxilliped; G, second maxilliped. Bar scale represents 0-5 mm. 



INDIAN OCEAN RANINID AND HOMOLID LARVAE 7 

Ocean raninids (see Table i) are completely unknown. The 1. 1. O.K. collections 
contain only four raninid zoeae and a single megalopa. One of these zoeae appar- 
ently belongs to R. ranina but the identities of the other specimens are very un- 
certain. 

The British Museum (Natural History) collections contain hatched material of 
R. ranina and I have taken this opportunity to re-illustrate the first zoea of this 
species. 

Ranina ranina (L.) 

(figs i and 2) 
Aikawa, 1941, pp. 117-118, fig. i. 

MATERIAL: (a) About 100 stage i zoeae hatched in the Aquarium de Noumea, 
New Caledonia, 4. XII. 1956 and presented to the British Museum (Natural History) 
by Mons. P. Budker of the Museum National d'Histoire Naturelle, Paris (B.M. reg. 
no. 1958: 7: 4: 2-9). 

(b) One stage i zoea taken by the Anton Bruun at station 18, 07 4i'N: 97 59 'E 
on 21. III. 1963. This specimen is badly damaged but appears to belong to this 
species. 

SIZE: Tip to tip of the rostral and dorsal carapace spines 6-7-7-7 mm - 

REMARKS: Because of the relative abundance of material available it is now 
possible to illustrate the larvae of this species adequately and a written description is 
not necessary. 

Aikawa's Japanese larvae were somewhat smaller than those from Noumea and 
he gives a spine tip to spine tip length of 4-2 mm, but the two sets of larvae show 
very close agreement in all other features which Aikawa either described or illus- 
trated. 

However, Aikawa did not mention the spinules on the antennae and maxillipeds 
although they were almost certainly present in his material. Williamson (1965) 
reported such spines on the maxillipeds of first stage zoeae of Lyreidus tridentatus, 
and these features may therefore be widespread within the Raninidae. However, 
without dissection the appendages of the larvae reported below could not be exam- 
ined at sufficiently high magnifications in all cases to determine whether the spinules 
are present or not. 

Raninid larva A; fRaninoides sp. 

(fig- 3) 

MATERIAL: One specimen in the first zoeal stage. Position 12 36'N: 8o4o'E; 
depth 200 m to surface. Date 20. VI. 1964. Vessel I.N.S. Kistna; Station 378. 
I.O.B.C. serial no. 0612. 

SIZE: Tip to tip of the rostral and dorsal carapace spines 3-3 mm; Carapace 
length from between eyes to postero-lateral carapace margin 0-93 mm. 



A. L. RICE 




FIG. 3. Raninid larva A, stage I zoea. A, Frontal view; B, lateral view; C, dorsal view of 
abdomen; D, antenna; E, first maxilliped ; F, second maxilliped. The setae covering the 
surface of the carapace and abdomen are omitted from B and C for clarity. Bar scales repre- 
sent i-o mm 



INDIAN OCEAN RANINID AND HOMOLID LARVAE g 

DESCRIPTION: Carapace with prominent, curved, dorsal and rostral carapace 
spines, each with a number of subsidiary spines, mainly on the anterior edge of the 
rostral spine and the posterior edge of the dorsal spine. Lateral spines normally 
directed downward and forward, roughly parallel to the rostral spine, but that of 
the left-hand-side displaced backwards (see fig. 3). Each lateral spine with a row of 
subsidiary spines on the dorsal edge. Anterior dorsal tubercle in the mid-line 
between eyes; posterior tubercle behind the dorsal spine. A pair of spines on each 
side of the posterior carapace tubercle, close to the posterior carapace margin. 
Surface of carapace covered with stiff, almost spine-like bristles (not shown in fig. 
3(B)). Eyes sessile, each with a proment spine and a papilla on the stalk. 

Abdomen of 5 segments plus the telson. Segment i with a large mid-dorsal spine 
and 2 pairs of smaller dorso-lateral spines. Segments 2-5 each with a single median 
ventral spine and paired dorsal, dorso-lateral, postero-lateral and postero- ventral 
spines. The dorso-lateral spines on segment 2 directed forwards like the 'lateral 
knobs' on this segment in the larvae of the higher Brachyura. 

Telson (fig. 3(C)). Each arm of the shallow telson fork with a very large setose 
spine and 3 postero-median plumed setae. Outside the major spine on each side 
there are 2 spines, the outer naked, the inner setose. Antero-lateral margin of each 
telson arm carries 4 smooth spines. There are a number of spinules on the dorsal 
surface of the telson, and each of the 2 major spines has a subsidiary spine on the 
dorsal surface close to the base. 

Antennule simple and unsegmented, with 2-3 terminal aesthetascs and i seta. 

Antenna as shown in fig. 3 (D). 

Mandibles with incisor and molar processes, but no palp. 

Maxillule with an unsegmented endopod with 3 terminal and i sub-terminal seta. 
Endites well developed, but no lateral seta on basis. 

Maxilla with 5 setae on the endopod. Scaphognathite with 4 sub-equal plumose 
setae on the lateral margin and a much longer posterior seta. 

Maxillipeds i and 2 as shown in fig. 3 (E and F). 

None of the more posterior appendages developed. 

REMARKS: This larva is tentatively attributed to the genus Raninoides, on the 
basis of its close resemblance to that of Raninoides benedicti, but it could belong 
to other Indian Ocean genera whose larvae are unknown. It differs so much from 
the first zoea of Lyreidus tridentatus, particularly in the length of the lateral carapace 
spines and in the form of the telson, that it is very unlikely to belong to L. channeri, 
the only other member of the genus recorded from the Indian Ocean. The larva also 
differs from the Ranina ranina larvae illustrated above, particularly in being much 
smaller and possessing eyestalk spines. 

Of the raninids recorded from the Indian Ocean this leaves 7 species belonging to 
the genera Cosmonotus, Notopus, Notosceles and Raninoides as possible parents. 
The larva could belong to any of these genera, but its very close resemblance to the 
described larvae of Raninoides benedicti Rathbun (Knight, 1968) makes another 
species of this genus a strong possiblity. 

The larva differs from the first stage of R. benedicti in only a few relatively minor 
points. Thus, the lateral carapace spines are relatively longer than in R. benedicti 



A. L. RICE 



B 




E 



FIG. 4. Raninid larva B, stage III (?) zoea. A, Frontal view; B, lateral view; C, dorsal view 
of abdomen; D, antenna; E, first maxilliped; F, second maxilliped. Bar scale represents 
i-o mm for D and 2-0 mm for the remainder. 



INDIAN OCEAN RANINID AND HOMOLID LARVAE n 

and carry more subsidiary spines. There are 4 spines on the antero-lateral margin 
of each telson fork as opposed to 2 in R. benedicti, and there is an extra pair of spines 
on the posterior carapace margin. All of these differences are probably specific 
rather than generic. 

If the larva does belong to Raninoides then the known distributions of the adults 
(Table i) indicate that it probably belongs either to R. personatus White or to 
R. serratifrons Henderson. 



Raninid larva B 

(fig. 4) 

MATERIAL: One slightly damaged specimen, probably a third zoea. Position 
06 26 'N: 49 46'E; depth 200 m to surface. Date 17. VIII. 1964. Vessel Argo; 
Dodo cruise; Station 37. I.O.B.C. serial no. 0374. 

SIZE: Carapace length from between the eyes to posterior carapace margin 
1-83 mm; Abdomen length 2-71 mm; no spine tip to spine tip measurement can be 
given because the dorsal spine is broken. 

DESCRIPTION: Carapace with long dorsal and rostral spines, and curved, for- 
wardly directed lateral spines, each with a number of subsidiary spines. Anterior 
and posterior blunt dorsal tubercles in the mid-line. A supra-ocular spine present 
on the right-hand-side; the left-hand-side is damaged, but does not appear to have 
possessed a spine in this position. A number of spines on the carapace surface 
beneath the insertion of each lateral spine and on the slightly raised ridge between 
the dorsal and lateral spines. A series of spines on each postero-lateral carapace 
margin decreasing in size ventrally. Eyes free, with the stalks carrying many 
setae but no sign of papillae or spines. Whole surface of carapace covered with 
short setae (not shown in the illustrations). 

Abdomen of 6 segments plus the telson. Segments 2-5 with long, acute spines on 
the postero-lateral corners. Segment 2 with forwardly directed lateral processes 
and segments 4 and 5 each with a median ventral spine. All abdominal segments 
with a number of other spines on the dorsal and lateral surfaces and also on the 
posterior margins; segments 1-5 have long setae on the posterior margins (fig. 4(C)). 

Telson (fig. 4(C)) a broad flat plate, about 4 times as broad as its length in the 
mid-line. Posterior margin on the right-hand-side carries a large, fused spine, 
presumably representing the 4th telson process. Outside this large spine there are 
a very small and 2 larger fused spines which probably represent the first 3 telson 
processes, although the outer one is not clearly distinguishable from the series of 
spines on the antero-lateral telson margin. Postero-lateral region of the telson on 
the left-hand-side is damaged beyond the base of the main fused spine. Posterior 
margin with 21 articulated processes between the major spines. A series of small 
spines on the dorsal surface, close to the posterior margin, extends onto the main 
telson spines. Dorsal surface of telson and abdominal segments with many small 
setae. 



12 



A. L. RICE 



Antennule simple and unsegmented, with 3-4 terminal aesthetascs and a single 
seta. 

Antenna (fig. 4(D)) with 19-20 marginal setae on the scale and a series of small 
bristles on both the upper and lower surfaces. Endopod less than 1/3 length of 




FIG. 5. Raninid larva C, terminal zoea. A, Dorsal view; B, lateral view of thorax; C, lateral 
view of abdomen; D, antenna; E, ventral view of last pleopod and uropod on the right-hand- 
side. The small surface setae have been omitted partly from A and wholly from B. Bar 
scales represent i-o mm. 



INDIAN OCEAN RANINID AND HOMOLID LARVAE 13 

scale, fused to basis and armed with a single terminal seta. Basis with spinous 
process about 1/3 length of scale, and a second prominent spine at the base of the 
scale. 

Manbidle with no palp. 

Maxillule with an unsegmented endopod armed with 4 setae. A single seta on 
the lateral margin of the basis below the endopod. 

Maxillipeds i and 2 as shown in fig. 4(E and F). Exopod of maxilliped i with n 
or 12 natatory setae, that of maxilliped 2 with 13 setae on the distal half and 3 on 
the proximal half. 

Maxilliped 3 and the more posterior thoracic appendages present as unarmed, 
unsegmented buds. 

No pleopods present but the uropods have well developed exopods carrying 6 or 7 
marginal setae. Endopods represented by very small unarmed buds not separated 
from the protopods. 

REMARKS : From the known development of Lyreidus tridentatus and Raninoides 
benedicti it is clear that the determination of the zoeal stage in raninid larvae is not 
as simple as it is, for example, in the higher Brachyura. However, from the pres- 
ence in the above larva of well developed uropods but the absence of pleopods, and 
also the degree of development of the antennules, antennae and maxillipeds, it 
appears to be at about the mid-point of the zoeal series, perhaps the third or fourth 
of a total of about 6 zoeal stages. 

Identification of this larva is also difficult. The differences between it and the 
known larvae of Lyreidus and Raninoides suggest that it belongs to a genus other 
than these. The changes which would be necessary to transform the first zoea of 
Ranina ranina into this larva, particularly in the armature of the carapace and the 
form of the telson, are greater than those ocurring during the development of L. 
tridentatus and R. benedicti, indicating that it does not belong to R, ranina either. 
This leaves the parentage still very uncertain, but probably among the genera 
Cosmonotus, Notopus and Notosceles. 

Raninid larva C 

(fig- 5) 

MATERIAL: One specimen in the last zoeal stage. Position 09 34'N: 75 i6'E; 
depth 200 m to surface. Date 10. II. 1965. Vessel Meteor; Station 187. I.O.B.C. 
serial no. 0145. 

SIZE: Tip of rostral spine to tip of dorsal spine 3-8 mm; tip of rostral spine to 
posterior median carapace margin 3-9 mm; total length (tip of rostral spine to base 
of telson fork) 6-6 mm. 

DESCRIPTION: Carapace with prominent forwardly directed rostral and paired 
lateral spines; subsidiary spines on the dorsal surface of the rostrum and on the 
outer surfaces of the laterals. Dorsal spine small, curving posteriorly. Blunt 
anterior and posterior carapace tubercles in the mid-line. Rostrum widens between 
the eyes to a front carrying a series of spines on each antero-lateral angle. A pair 



14 A. L. RICE 

of bulbous lobes behind the eyes each carrying a prominent spine antero-laterally 
and a number of smaller spines dorsally. A curving row of 6 spines on each side 
of the carapace between the bases of the lateral and dorsal spines. A parallel row of 
4 spines close to the postero-lateral carapace margin. Each postero-ventral corner 
of the carapace with a single spine and a series of denticles. A prominent papilla on 
each eyestalk. 

Abdomen (fig. 5 (A and C)) of 6 segments plus the telson. First segment with a 
raised transverse ridge armed with a row of long plumose setae and with 3 spines 
close to the mid-line. Segments 2-5 all basically similar, having each postero- 
lateral margin produced into a long, slightly curved spine. A somewhat variable 
number of other spines on each segment, but a basic pattern of 4 on the dorsal 
surface anteriorly, and a series of 5 on the postero-dorsal margin. Segment 5 with 
additional spines, including a prominent one at each postero-lateral corner. Seg- 
ment 6 with only 2 dorsal spines, but 6 spines on the posterior margin, the outer 
pair in dorsal view looking like the postero-lateral spines of the more anterior seg- 
ments. 

Telson (fig. 5 (A)) a broad, bilobed plate. Each antero- and postero-lateral margin 
with a series of fused spines increasing in size gradually posteriorly, but ending in a 
much larger spine with a small spine basally on the dorsal surface. Posterior telson 
margin with 10 pairs of articulated processes between the telson forks, inner 3 pairs 
much smaller than the others and probably added at the most recent moult. 

The whole of the dorsal surface of the carapace, abdomen and telson covered with 
short, close-set setae. 

Antennule with an unsegmented peduncle, swollen basally and with an obvious 
statocyst. Endopod unarmed and not separated from the peduncle. Flagellum 
with 3 groups of about 3, 3, and 4 aesthetascs. 

Antenna (fig. 5(D)) with 26-28 marginal setae on the scale. Endopod 3-segmented, 
about as long as scale and with a single terminal seta. Protopod with 2 spines and 

3 setae. 

Mandible with an unarmed, unsegmented palp. 

Maxillule with 2-segmented endopod carrying 5 setae on the distal segment and a 
single seta on the proximal segment. Lateral margin of basis with a single seta. 

Maxilla with more than 70 marginal setae on the scaphognathite. Unsegmented 
endopod with 6 terminal, i medial and 4 lateral setae. 

Maxilliped i with 2-lobed epipod. Basipod with 15 setae on medial margin. 
The 5 segments of the endopod carry 3, 2, 2, 4 and 6 setae respectively on the medial 
margins. The lateral margins of segments 2 and 5 (terminal) each with a single 
fine seta, those of segments 3 and 4 each with 2 fine setae. Exopod 2-segmented; 
proximal segment with 8 setae along posterior edge, distal segment with 15 or 17 
setae around whole margin. 

Maxilliped 2 with simple epipod. Basipod with 4 setae on medial margin and 2 on 
posterior surface at the base of the endopod and exopod. Endopod on one side of 

4 segments with 2, i, 2 and 3 inner setae respectively. On the other side the long 
penultimate segment is sub-divided, the proximal part being unarmed. Terminal 
and sub-terminal segments each with a very fine lateral seta. Exopod 2-segmented ; 



INDIAN OCEAN RANINID AND HOMOLID LARVAE 15 

proximal segment with 6 or 7 setae along posterior edge, distal segment with 21 
marginal setae. 

Maxilliped 3 with endopod indistinctly divided into 4 segments of which the 
terminal and sub-terminal each carry a short seta. Exopod short, simple and 
unarmed. 

Pereiopods all present and indistinctly segmented, the first pair chelate. 

Pleopods on abdominal segments 2-5 well developed but unsegmented, with small 
endopods and larger exopods (fig. 5 (C and E)). Exopods and endopods unarmed, 
but with somewhat serrate margins indicating that they would probably become 
setose at the next moult. Uropods with exopods separated from protopods and 
carrying 20-21 plumose setae. Endopods represented by small buds and not 
separated from the protopods (fig. 5(E)). 

REMARKS: Of the raninid larvae previously described, this zoea most closely 
resembles Lithozoea serrulata described by Aikawa (1933). The two larvae share 
a number of common features including the broad, plate-like telson, forwardly 
directed carapace spines, spine rows on the posterior carapace margin and on the 
side of the carapace between the lateral spines and the dorsal spine, and relatively 
short rostral and dorsal spines. The main differences between the larvae are the 
development of the anterior carapace tubercle into a bifurcated spine and the pres- 
ence of lateral carapace keels in Lithozoea, the presence of a mandibular palp in the 
Indian Ocean larva and 4 ot 5 segments in the endopod of the second maxilliped 
in this larva compared with only 3 in Lithozoea. 

Williamson (1965) pointed out that Lithozoea serrulata possesses a number of 
homolid characters, particularly in the carapace spines and keels. Although raninid 
larva C is somewhat less homolid in these respects than Lithozoea, it does resemble 
late homolid zoeae in having more than 3 segments in the endopod of the second 
maxilliped. No brachygnathan larvae have more than 3 segments in this endopod, 
and the only previously described raninid larva which may have more than 3 is 
Acanthocaris described by Claus (1876 and 1885) (see Williamson, 1965, p. 388), 
which also shows some homolid features in its carapace. 

Little can be said about the identity of this Indian Ocean larva. As with the 
previous larva, comparison with published larval descriptions readily excludes it 
from the genera Lyreidus, Ranina and Raninoides. This still leaves as possible 
parents the genera Notopus, Cosmonotus and Notosceles, although adult Notosceles 
are so similar to Raninoides that they must surely have similar larvae. The simil- 
arities between the Indian Ocean larva and Aikawa's Lithozoea serrulata suggest that 
they belong to the same genus. If this is so, the Indian Ocean larva may belong to 
Notopus dorsipes (Fabr.) since, of the three genera deduced as possible parents, 
Notopus is the only one with different species recorded in Indian and Japanese 
waters. N. dorsipes is the only species known from the Indian Ocean, having been 
recorded by Alcock (1896) from off the Malabar coast, relatively close to where 
raninid larva C was collected, but two other species, N. ovalis Henderson and N. 
misakiensis Sakai are also recorded from Japan (Sakai, 1937). Lithozoea serrulata 
perhaps belongs to one of these species. 



i6 



A. L. RICE 



Raninid larva D; fRaninoides sp. 

(figs 6 and 7) 

MATERIAL: One megalopa. Position 10 36'N: 95 39'E; depth 125 to 250 m. 
Date 25. III. 1963. Vessel Anton Bruun; cruise i; Station 24 S.O.S.C. Ace. no. 3. 

SIZE : Carapace length from tip of rostrum to posterior carapace margin 3 -6 mm ; 
Maximum carapace width 1-9 mm. 

DESCRIPTION: The megalopa is fully illustrated in figs 6 and 7. It is very 
similar to the megalopa of Raninoides benedicti as described by Knight (1968) and 
therefore only the main differences between the two larvae will be mentioned here. 




G 



FIG. 6. Raninid larva D; ? Raninoides megalopa. A, Dorsal view; B, lateral view of cara- 
pace; C, ventral view of anterior part of thorax; D, dorsal view of telson; E, cheliped, left- 
hand-side, ventral view; F, G, and H, third, fourth and fifth pereiopods respectively of the 
right-hand-side, dorsal views. Bar scales represent i-o mm for A, B, C and E, and 0-5 mm 
for D, F, G and H. 



INDIAN OCEAN RANINID AND HOMOLID LARVAE 




FIG. 7. Raninid larva D ; ?Raninoides megalopa. A, Maxillule; B, maxilla; C, D and E, first, 
second and third maxillipeds in ventral view; F, ischium and exopod of third maxilliped in 
dorsal view. Bar scale represents i-o mm. 



i8 A. L. RICE 

The carapace is slightly narrower posteriorly than is that of R. benedicti. The 
eyestalk papillae are placed postero-ventrally rather than antero-dorsally as in R. 
benedicti. The large spine between the bases of the chelipeds in R. benedicti is 
absent. The Indian Ocean megalopa has no podobranch on the second maxilliped, 
while the ischium of the third maxilliped (fig. y(E and F)) is relatively longer than in 
R. benedicti and has no teeth on the inner margin. 

The chelipeds and the second and third legs in R. benedicti each have a prominent 
spine on the ventral margin of the merus; the Indian Ocean megalopa lacks these 
spines but has a spine on the dorsal margin of the carpus of legs two and three. 
(These two legs are very similar, and therefore only leg three is illustrated in fig. 6.) 
The fourth and fifth legs are very similar in shape in the two larvae, but the fourth 
leg in the Indian Ocean specimen carries many more setae and the dactyl is less 
acutely pointed than in R. benedicti. 

REMARKS: The similarity between this larva and the known megalopa of Ran- 
inoides benedicti indicates that it belongs to the same genus. From the locality 
of capture of the larva and the known distributions of the adults, Raninoides hender- 
soni Chopra and R. serratifrons Henderson are equally likely to be the parent. 

Family HOMOLIDAE 

Plankton-caught larvae of the Homolidae present very much the same problems 
of identification as do those of the Raninidae, since larvae have been hatched from 
only four species and the complete larval development is known for only one of these 
(see Williamson, 1965 and Rice and Provenzano, 1970). 

The 1. 1. O.K. collections contain only two homolid zoeae, neither of them agreeing 
with any previously described larvae. 

Homolid larva A fHomola sp. 

(fig. 8) 

MATERIAL: One specimen, probably a second zoea. Position 11 49 'S: 49 23 'E. 
Date 21. VII. 1964. Vessel R.R.S. Discovery; Station 5508. I.O.B.C. serial no. 



SIZE : Carapace length from tip of rostrum to posterior margin in mid-line i -4 mm ; 
Total length from tip of rostrum to posterior margin of telson in mid-line 2-5 mm. 

DESCRIPTION : The main features of the morphology of this larva are adequately 
shown in the illustrations. This description will therefore be restricted to those 
features which are not illustrated but which can be seen without dissection. 

Antennule 2-segmented. Distal segment with 3 or 4 aesthetascs and setae; 
proximal segment with a long median seta, representing the inner flagellum, and a 
row of 3 short setae. 

Mandibles without palps. 

Maxilla with about 18 marginal setae on the scaphognathite. 

Maxilliped I as shown in fig. 8 (D). 



INDIAN OCEAN RANINID AND HOMOLID LARVAE 19 

Maxilliped 2 with the 4 segments of the endopod carrying i, i, 2 and 5 setae 
respectively, one seta on the terminal segment being laterally placed. Basis with 5 
medial setae and the exopod with 7 natatory setae. 

Maxilliped: j with an unsegmented endopod carrying a single terminal seta. 
Exopod with 7 setae. 

Posterior thoracic appendages represented by unarmed, unsegmented buds. 




D 



FIG. 8. Homolid larva A, stage II zoea. A, Dorsal view of carapace; B, dorsal view of 
abdomen; C, lateral view; D, antenna; E, first maxilliped. Bar scale represents i-o mm. 



20 A. L. RICE 

REMARKS: In general form this larva is very similar to the larvae of Homola 
barbata (Fabr.) described from Florida by Rice and Provenzano (1970), and also 
to the series of larvae taken in the plankton off South Africa and attributed to a 
species of Homola by Rice and von Levetzow (1967). However, the present specimen 
differs from these previously described forms in several important respects. 

The supra-ocular spines and the anterior and posterior carapace tubercles, which 
are very prominent in the Florida and South African larvae, are much reduced in the 
Indian Ocean specimen. This larva also has the anterior end of the dorsal tooth 
row on the carapace less protruberant than in the other larvae, and less likely to be 
replaced by a spine in the later stages. 

The dorsal spines on abdominal segments 2-5 in the Florida and South African 
larvae are entirely lacking in this specimen and, except for the second segment, the 
dorso-lateral spines are reduced to blunt protruberances. The Indian Ocean larva 
also lacks spines on the dorsal surface of the telson. 

Finally, in its degree of development this larva does not agree precisely with any 
stage in either of the previously described series, falling between the second and 
third stage in both cases. 

The absence of dorsal telson spines in two 1 Homola larvae from off south-east 
Africa described by Boas (1880) suggested to Rice and von Levetzow that these 
larvae were specifically distinct from their own South African material. From the 
known distributions of the adults these authors suggested that Boas's larvae might 
represent Homola orientalis Henderson, while their own material belonged to the 
eastern Atlantic and South African form of H. barbata. 

The larvae described here also lacks dorsal telson spines, like Boas's larvae, but 
differs from them in having reduced supra-ocular spines. Boas's larvae also 
have a pair of dorso-lateral carapace spines, presumably developed from the anterior 
end of the dorsal carapace tooth row of an earlier stage ; as noted above, such spines 
are unlikely to be developed in a later stage of the I.I.O.E. larva. If these differences 
are specific, then there is either a third, unsuspected, species of Homola in the south- 
western Indian Ocean, or else at least some of the larvae attributed to Homola in 
fact belong to a different genus. 

Knowledge of generic differences between homolid larvae is restricted to a compari- 
son of hatched larvae of Homola barbata (Fabr.) (Rice and Provenzano, 1970), 
Paromola japonica Parisi and Latreillia phalangium de Haan (Aikawa, 1937) and 
Latreillia australiensis Henderson (Williamson, 1965). Williamson noted that the 
homolid larvae described up to that time (1965) fell into two groups, one of which, 
including the hatched larvae of Latreillia and Paromola, differed from the second 
group in having neither dorsal nor antero-lateral carapace spines and no dorsal 
spines on the abdomen. Williamson suggested that these groups probably did not 
represent taxonomic groupings within the family and that intermediate forms would 
probably be found as more homolid larvae were described. 

The larva described here agrees most closely with the second of these larval 
groups, which includes not only the larvae described by Rice and von Levetzow 
(1967), Rice and Provenzano (1970) and Boas (1880), but also Gurney's (1924) 
Dromiacean species I and specimens attributed to the genus Homola by Cano (1893), 



INDIAN OCEAN RANINID AND HOMOLID LARVAE 



Thiele (1905), Pike and Williamson (1960) and Rice (1964). However, it differs 
from all of these forms in the absence of mid-dorsal spines on the abdominal segments 
and is therefore intermediate between Williamson's two groups. Considering the 
great similarity between the known larvae of Latreillia and Paromola, generic 
differences between homolid larvae seem to be very slight so that this larva may, 
indeed, belong to a genus other than Homola. 






FIG. 9. Homolid larva B, stage III zoea. A, Dorsal view; B, lateral view; C, antenna; D, E 
and F, first, second and third maxillipeds. Bar scale represents i-o mm for A and B, and 
0-5 mm for C, D, E and F. 



22 A. L. RICE 

Homolid larva B 

(fig- 9) 

MATERIAL: One specimen, probably a third zoea. Position 16 3i'N: 54 o8'E. 
Date 30. VI. 1963. Vessel R.R.S. Discovery: Station 5026. I.O.B.C. serial no. 0811. 

SIZE : Carapace length from tip of rostrum to posterior margin in mid-line i -5 mm. 
Total length from tip of rostrum to posterior margin of telson in mid-line 
3 -3 mm. 

DESCRIPTION: Carapace with a forwardly directed and somewhat upturned 
rostrum. Prominent mid-dorsal spine and smaller posterior carapace tubercle; no 
anterior tubercle. Dorsal tooth row of 70-80 teeth on each side, extending from 
middle of posterior margin to behind the eye and ending in a rather longer and 
stouter tooth. Postero-ventral tooth row of 50-60 sub-equal teeth. A prominent 
ventro-lateral spine on each side of the carapace above the base of the third maxil- 
liped. Prominent eyestalk papillae. 

Abdomen of 6 segments and telson. Segment 2 with slight dorsal transverse ridge 
and segment 5 with small postero-lateral processes; otherwise all abdominal seg- 
ments simple and unarmed apart from some setae close to posterior margins. 

Telson a broad, triangular plate, with 3 fused spines at each postero-lateral angle 
and 10 pairs of processes articulated to posterior margin. Uropods with protopods 
not separated from endopods and exopods. Endopods unarmed, exopods with 
9-11 marginal setae. 

Antennule 2-segmented, proximal segment with a single long plumose seta repre- 
senting ventral flagellum, distal segment with 2 terminal aesthetascs and 2 setae. 

Antenna (fig. g(C)) with 16 marginal setae on scale, spinous process about 1-5 times 
length of scale, endopod unarmed and less than half length of scale. 
Mandibles without palps. 

Maxillule with 5 setae on distal segment of endopod and a single seta on proximal 
segment ; no lateral seta on basis. 

Maxilla with about 40 setae on scaphognathite. 

Maxillipeds as illustrated in figs 9 (D, E, and F) ; exopods of each with 8 natatory 
setae. 

Posterior thoracic appendages present as unarmed, unsegmented buds. 

REMARKS: Like the preceding larva, this specimen is intermediate between the 
two homolid larval groups noted by Williamson. Thus, although it possesses the 
prominent dorsal carapace spine of the 'Homola group, it lacks the dorsal abdom- 
inal spines typical of most of these larvae. It is even less Homola-like. than the larva 
described above since it also lacks any suggestion of supra-ocular spines, it has very 
small antero-laterals (if, indeed, the enlarged teeth at the anterior end of the dorsal 
carapace tooth rows represent these), and it has no dorso-lateral projections on the 
abdominal segments. The combination of characters is so different in this larva 
from that in any previously described zoea that its identity cannot be established at 
present. 



INDIAN OCEAN RANINID AND HOMOLID LARVAE 23 

DISCUSSION 

The small collection of raninid and homolid larvae reported here tends to confirm 
Williamson's (1965) conclusions that the similarities between the two groups are 
consistent with both families being fairly close to a pre-brachyuran stock, the 
raninids being considerably more 'brachyuran' than the homolids. 

Perhaps the most interesting larva in the collection is that described as raninid 
larva C which, like Aikawa's Lithozoea serrulata, is intermediate in many features 
between the homolids and the typical raninid larvae. Williamson noted that L. 
serrulata represents a possible intermediate step in the simplification of the rather 
complex carapace armature of the homolids towards the dorsal, rostral and lateral 
spines of the carapace in typical raninid and brachygnathan zoeae. Raninid larva C 
seems to be a further step in this direction, since the homolid denticulate carapace- 
folds, still present in L. serrulata, are represented only by a series of short spines in 
this larva. The relationship of the lateral carapace spines to the carapace folds and 
denticle rows in raninid larva C and L. serrulata indicate that the lateral spines in the 
raninids, and presumably also in the higher Brachyura, are homologous with either 
the antero-lateral spines in the homolids or, more probably, with the spines developed 
from the anterior ends of the dorsal denticle rows. 

The bilobed, plate-like telsons of L. serrulata and raninid larva C also conveniently 
bridge the gap between the rather anomuran broad triangular telsons of late homolid 
zoeae and the forked telsons of the zoeae of Ranina, Raninoides and Lyreidus, which 
are more similar to those of typical brachyuran zoeae. 

It would be interesting to know if these seemingly primitive raninid larvae belong 
to species with similarly primitive adults, but such information can come only from 
the rearing work which is much needed in both the Homolidae and the Raninidae. 

ACKNOWLEDGEMENTS 

My thanks are due to the Directors of the Indian Ocean Biological Centre and the 
Smithsonian Sorting Center for allowing me to examine material in their care. I 
would also like to thank Dr. K. K. Tiwari of the Zoological Survey of India and Dr. 
Henry B. Roberts of the Smithsonian Institution for providing me with details of 
adult raninids from the Indian Ocean in their collections. 

REFERENCES 

AIKAWA, H. 1933. On larval forms of some Brachyura. Paper II: a note on indeterminate 

zoeas. Rec. Oceanogr. Wks. Jap. 5 : 124-254. 
1937. Further notes on brachyuran larvae. Rec. Oceanogr. Wks. Jap. 9 : 87-162. 

- 1941. Additional notes on brachyuran larvae. Rec. Oceanogr. Wks. Jap. 12 : 117-120. 
ALCOCK, A. W. 1896. Materials for a carcinological fauna of India. No. 2. The Brachyura 

Oxystomata. /. Asiat. Soc. Beng. 65 (II) : 134-296. 

- 1899. An account of the deep-sea Brachyura collected by the Royal Indian Museum 
Maritime Survey Ship "Investigator". Calcutta, 85 pp. 

BARNARD, K. H. 1950. Descriptive catalogue of South African decapod Crustacea (Crabs and 

shrimps). Ann. S. Afr. Mus. 38 : 1-837. 
BIANCONI, G. G. 1851. Specimina Zoologica Mosambicana. Fasc. 5 : Memorie R. Accad. Sci. 

1st. Cl. Sci.fis. Bologna, 3 : 91-112. 



24 A. L. RICE 

BOAS, J. E. V. 1880. Studier over Decapodernes Slaegskabsforhokl. K. danske Vidensk. 

Selsk. Skr. (6) Nat. og Math., Afd. 1 (2) : 25-210. 
BOUVIER, E. L. 1915. Decapodes marcheurs (Reptantia) et Stomatopodes recueillis a 1'ile 

Maurice par M. Paul Carie. Bull, scient. Fr. Belg. (7) 48 (3) : 178-318. 
CANO, G. 1893. Svillupo del Dromidei. Atti Accad. Sci. fis. mat., Napoli 6 (2) : 1-23. 
CHOPRA, B. i933a. Further notes on Crustacea Decapoda in the Indian Museum. III. On 

the decapod Crustacea collected by the Bengal Pilot Service off the mouth of the River 

Hughli. Dromiacea and Oxystomata. Rec. Ind. Mus. 35 : 25-52. 
- !933b. Further notes on crustacean Decapoda in the Indian Museum. IV. On two new 

species of oxystomous crabs from the Bay of Bengal. Rec. Ind. Mus. 35 : 77-86. 
CLAUS, C. 1876. Untersuchungen zur Erforschung der genealogischen Grundlage des Crustaceen- 

Systems. Ein Beitrage zur Descendenzlehre. VVein, 114 pp. 
CLAUS, C. 1885. Neue Beitrage zur Morphologic der Crustaceen. Arb. zool. Inst. Wien, 6 : 

1-108. 

DOFLEIN, F. 1904. Brachyura. Ergebn. dt. Tiefsee-Expedn 'Valdivia' 1898-1899. 6 :i-3i4- 
GURNEY, R. 1924. Decapod larvae. Nat. Hist. Rep. Br. Antarct. Terra Nova Exped. (Zool.) 

8 : 37-202. 
HENDERSON, I. R. 1893. A contribution to Indian carcinology. Trans. Linn. Soc. Lond. (2) 

5 : 325-458. 
HOFFMANN, C. K. 1874. Crustaces et Ecinodermes. In: F.P.L. Pollen and D.C. Van Dam 

(Ed.) Recherches sur la faune de Madagascar 5 (2) : 1-58. 
KNIGHT, M. D. 1968. The larval development of Raninoides benedicti Rathbun (Brachyura, 

Raninidae), with notes on the Pacific records of Raninoides laevis (Latreille). Crustaceana, 

Suppl. 2 : 145-169. 
LAURIE, R. D. 1906. Report on the Brachyura collected by Prof. Herdman at Ceylon 1902. 

Rep. Pearl Oyster Fish. 5, Suppl. Rep. XL: 349-432. 
NOBILI, G. 1905. Crostacei di Zanzibar. Boll. Mus. Zool. Anat. comp. R. Univ. Torino. 20 

(506) : 1-12. 
PIKE, R. B. and WILLIAMSON, D. I. 1960. Larvae of decapod Crustacea of the families 

Dromiidae and Homolidae from the Bay of Naples. Pubbl. Staz. zool. Napoli 31 : 553-563. 
RICE, A. L. 1964. The metamorphosis of a species of Homola (Crustacea, Decapoda: Drom- 
iacea). Bull. mar. Sci. Gulf Caribb. 14 : 221-238. 

and PROVENZANO, A. J. 1970. The larval stages of Homola barbata (Fabricius) (Crus- 
tacea, Decapoda, Homolidae) reared in the laboratory. Bull. mar. Sci. (20 : 446-471). 

and VON LEVETZOW, K. G. 1,967. Larvae of Homola (Crustacea, Dromiacea) from South 
Africa. /. nat. Hist. 1 : 435-453. 

SAKAI, T. 1937. Studies on the crabs of Japan. II. Oxystomata. Scient. Rep. Tokyo 

Bunrika Daigaku Sect. B 3, Suppl. no. 2 : 67-192. 
THIELE, J. 1905. Uber einige stielaugige Krebse von Messina. Zool. Jb., Suppl. 8 (Festschr. 

f. Mobius) : 443-474. 
WARD, M. 1942. A new genus and eight new species of Brachyura from Mauritius and the 

Chagos Archipelago. Bull. Maurit. Inst. 11 (2) : 39-48. 
WILLIAMSON, D. I. 1965. Some larval stages of three Australian crabs belonging to the 

families Homolidae and Raninidae, and observations on the affinities of these families 

(Crustacea: Decapoda). Aust. J. Mar. Freshw. Res. 16 : 369-398. 
WOOD-MASON, J. 1886. Description of a new species of Crustacea belonging to the brachy- 

urous family Raninidae. /. Asiat. Soc. Beng. 56 : 206-209. 

Dr. A. L. RICE, 

Department of Zoology 

BRITISH MUSEUM (NATURAL HISTORY) 

CROMWELL ROAD 

LONDON, S.W.7 




Printed in England by Staples Printers Limited at their Kettering, Northants, establishment 



BATS FROM ETHIOPIA COLLECTED 
BY THE GREAT ABBAI EXPEDITION, 

1968 



J. E. HILL 

AND 

P. MORRIS 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 21 No. 2 

LONDON: 1971 



<? 



I 2 7 APR 1971 

BATS FROM ETHIOPIA COLLECTED BY 
THE GREAT ABBAI EXPEDITION, 1968 




BY 

JOHN EDWARDS HILL 

AND 

K 
PATRICK MORRIS 



Pp. 25-49; 3 Plates 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 21 No. 2 

LONDON : 1971 



THE BULLETIN OF THE BRITISH MUSEUM 

(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 21, No. 2 of the Zoological series. 
The abbreviated titles of periodicals cited follow those 
of the World List of Scientific Periodicals. 



World List abbreviation 
Bull. Br. Mus. not. Hist. (Zool.) 



Trustees of the British Museum (Natural History), 1971 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 28 April, 1971 Price 1-15 



BATS FROM ETHIOPIA COLLECTED BY 
THE GREAT ABBAI EXPEDITION, 1968 

By J. E. HILL & P. MORRIS 

INTRODUCTION 

THERE has been hitherto no great concentration on the bat fauna of Ethiopia by 
collectors and many records are of one or two specimens sporadically obtained. 
For this reason the present collection is of particular interest, and is of especial 
value for the wide representation of the bats of western Ethiopia, hitherto largely 
unrecorded. This paper is an account of the bats collected by the Great Abbai 
Expedition including taxonomic notes, and revisions where appropriate, together 
with what little ecological information is available. 

The principal aim of the Expedition was to carry out a survey of the Blue Nile 
( Great Abbai) Gorge. The organization of the Expedition and its main activities 
are described by Blashford Snell (1970). The bat collections were made mainly in 
the Gorge itself at river level at various "Forward Bases" west of Shafartak Bridge, 
or in the Awash National Park, with some additional specimens being obtained 
elsewhere, notably in the highlands at Ghimbi. 

The Nile Gorge is a unique habitat, mostly uninhabited with dense scrub covering 
the steep rocky hillsides. There is a spectacular seasonal change here; during the 
dry season the river almost ceases to flow and the Gorge becomes exceedingly arid 
with only a narrow fringe of tall trees beside the river retaining their leaves. At 
this time survival must be a serious problem for many animals, Megachiroptera 
especially being unlikely to find any succulent fruits and presumably migrating 
elsewhere. By contrast insects appear to be especially abundant in the dry season, 
according to previous explorers, so that the Microchiroptera may well remain 
throughout the year, though the fierce heat and lack of shelter must cause problems. 
The Expedition's visit to the Nile Gorge came towards the end of the wet season, 
when the atmosphere was warm and humid; the lush green vegetation seemed to 
provide an ideal habitat for a wide range of animals and gave no indication of just 
how inhospitable the habitat is during at least one third of the year. 

The Awash National Park is chiefly an open dusty area of grassland, thorn scrub 
and volcanic rubble. An important geological feature, so far as bats are concerned, 
is the occurrence of "lava blister caves". These are evidently formed by gas 
bubbles in flowing lava and appear on the ground as low, rounded hillocks standing 
up prominently in an otherwise flat area. These mounds are hollow inside and 
some have perforated so allowing access to the interior. They represent the only 
cool, shady retreats in an otherwise open, arid landscape and consequently house 
many bats. Two of the blisters had openings only at their highest points. These 
had functioned as pitfall traps and also perhaps as carnivore dens, and the floors of 
both were littered with mammal bones which included the remains of five bat 
species. 



28 J. E. HILL & P. MORRIS 

Live bats were collected either by shooting (with -22 or -410 guns loaded with 
dust shot) or with mist nets. Fixed nets, supported on poles, trees or convenient 
rocks were avoided by flying bats, and only proved useful when set at a roost entrance 
whence the bats could be driven into the net. For catching bats as they flew 
low hawking for food a hand held net was employed. A small net (about i m x 
1-5 m), supported between two fibreglass rods was held close to the ground and 
flicked sharply upwards to catch the bat as it flew past. The method requires both 
patience and a certain amount of practice and was used with particular success by 
Dr. D. W. Yalden to catch most ol the free-flying bats obtained. Specimens taken 
this way come in ones and twos and contrast with sizeable batches of bats taken at 
roosts. Because roosting bats are easier to get in quantity, they figure prominently 
in this collection, though their numbers here are not necessarily a reflection of true 
abundance in the field. Netted bats are often of greater interest purely because 
they are difficult to catch, particularly if they do not form communal roosts ; they 
are therefore poorly represented in collections. 

The specimens obtained were either preserved whole in formalin, supported on 
card so as to keep the wings well displayed or they were prepared as a skull and dry 
card-mounted skin. The collections of the British Museum (Natural History) 
contain also a small number of specimens representing species previously not recorded 
from Ethiopia and notes on these are included in this report. Such specimens are 
denoted by their registration numbers, as is a single specimen from French Somali- 
land (Territory Afars and Issas) which constitutes a new record for that country and 
is included in this paper. 

The majority of bat specimens collected by the Great Abbai Expedition are now 
in the collections of the British Museum (Natural History), but where numbers have 
permitted this to be done, duplicate examples have been sent to the Museum at 
Haile Selassie I University, Addis Ababa. 

In the present paper, the altitudes and co-ordinates used have been taken from 
the best available maps, the i = 1,000,000 series. For convenience, clarity and 
ease of future reference these have been adhered to, even though investigations in 
the field showed the maps to be inaccurate in places. Part of the Nile forms the 
boundary between the provinces of Go j jam and Wollega; specimens obtained at 
the riverside have not been specifically assigned to either. Times given are 'local 
time', and as a rough rule the period 19.00 hrs-o6.oo hrs is passed in darkness. 
Linear measurements of specimens are in millimetres : the minimum, maximum and 
mean (in parentheses) are given for series. 

It is hoped that a future paper will give detailed background information on the 
ecology of the areas visited by the Expedition, meanwhile the brief survey above 
and the "collection and field notes" for the various species below have been prepared 
by one of us (P.M.) from actual field data; determinations and taxonomic investiga- 
tions are the work of J.E.H. We would like to express our thanks to various 
members of the Great Abbai Expedition for the hard work and long hours spent 
in pursuit of bats under trying conditions. The efforts in the field of Drs. D. W. 
Yalden, M. J. Largen and Mr. H. King are particularly acknowledged. 



BATS FROM ETHIOPIA 29 

SYSTEMATIC SECTION 

Epomophorus anurus (Heuglin, 1864) 

SPECIMENS. 2 young adult females. Temporary base, Mouth of Azir River, 
Blue Nile Gorge, 10 29' N, 36 25' E, alt. 1,000 m. 21 August 1968. 

TAXONOMIC NOTES. These young adult specimens agree closely in palatal propor- 
tions with those given by Anderson (1912 : 533) for females of E. anurus. Kock 
(1969 : 18) considers anurus a subspecies of E. labiatus. 

COLLECTION AND FIELD NOTES. A small group of fruit bats was located, flying 
about under the big trees lining the river; the bats were particularly concentrated 
around some large fig trees which bore plenty of ripe fruit. The remains of many 
chewed figs littered the ground below. The bats were seen at about 21.30 hours, 
and were immediately recognized as Megachiroptera by the distinctive way in which 
their eyes appeared large and red as they reflected the light from torch beams. Three 
specimens were obtained using dust shot. It was interesting to note a form of 
communal behaviour ; the bats had been following each other in threes and fours 
through gaps in the foliage, and when each specimen was shot, the others changed 
course and flew low and close to investigate the victim. The bats showed particular 
concern in response to the cries of a wounded individual. No other instances of 
group behaviour of this nature were observed with any of the other bats obtained. 
This communal response is all the more remarkable if the group really did include 
two species (see below). All three specimens of Epomophorus were females, so it 
may be that the group was part of a nursing colony whose individuals would perhaps 
have a stronger attachment to others of their species than normal. 

Epomophorus sp. 

SPECIMEN, i young adult female. Temporary Base, mouth of Azir River, Blue 
Nile Gorge, 10 29' N, 36 25' E, alt. 1,000 m. 21 August 1968. 

TAXONOMIC NOTES. It seems likely that this young adult specimen is referable 
to E. gambianus, a species recorded from southern Ethiopia by Andersen (1912 : 
540). Although from the proportions of the palate it could be referred to 
E. crypturus as defined by that author, this species is known so far only from localities 
in and south of the southern Congo and southern Tanzania, and, furthermore, in 
several of its dimensions this specimen exceeds the greatest size as yet recorded for 
E. crypturus. Measurements: length of forearm 77-9; greatest length of skull 50-3; 
condylobasal length 50-0; condylocanine length 48-3; median palatal length 27-7; 
post palatal length 12-3; rostral length 19-7; zygomatic width - ; least inter- 
orbital width 7-5; post orbital width 9-4; width of braincase 16-8; mastoid width 
17-2; ci-c 1 (alveoli) 8-9; mi-m 1 13-6; c-m 1 18-2; length of mandible 39-3; 
c-m 2 19-6. 

COLLECTION AND FIELD NOTES. As for E. anurus. If the present specimen really 
is a different species, it is surprising that it formed part of such a closely knit 
social group with E. anurus. 



30 J. E. HILL & P. MORRIS 

Micropteropus pusillus (Peters, 1868) 

SPECIMENS. Two females and a foetus. Sirba, Blue Nile Gorge, ioo5'N, 
35 30' E, alt. 800 m. 30 August 1968. 

TAXONOMIC NOTES. There appears to have been hitherto no confirmed record of 
Micropteropus from Ethiopia, although M. pusillus occurs in the southern Sudan 
(Kock, 1969 : 24). 

COLLECTION AND FIELD NOTES. Found at night between 20.00 hrs and 21.00 hrs 
flying around under large fig trees at the edge of the Nile. Both specimens caught 
in small hand-held mist nets. One animal was found to be pregnant with a well 
formed foetus, suggesting that breeding may coincide with the wet season (just 
ending at this time) and maximum availability of fruit such as figs. 

ADDITIONAL MATERIAL. Apart from the specimens obtained by the Great 
Abbai Expedition, the collections of the British Museum (Natural History) include 
three further Ethiopian specimens (B.M. 28.1.11.5-7) obtained at the Donkam 
River, Great Abbai, 100 miles southwest of Lake Tana, at 5,000 feet, by 
R. E. Cheesman, which presumably led Ellerman, Morrison-Scott and Hayman 
(1953 : 49) to include Ethiopia in the distribution of the species. 



Rhinopoma hardwickei sennaarinese Fitzinger, 1866 
(Plate 3 (a)) 

SPECIMENS, (i) Three males, three females. North eastern slope of Mount 
Fantalle, Shoa. 08 58' N, 39 54' E, alt. 1,000 m. 28 September 1968. 

(2) Two males, nine females. North bank of Awash River, Awash National 
Park, Shoa. 08 30' N, 40 01' E, alt. 1,000 m. 25-28 September 1968. 

(3) Skulls from cave deposit. Near Metahara, Awash National Park, Shoa. 
08 50' N, 40 01' E, alt. 1,000 m. 28 September 1968. 

TAXONOMIC NOTES. Kock (1969 : 35) has reviewed R. hardwickei with particular 
reference to forms inhabiting northeastern Africa and has concluded that larger 
specimens from the Sudan, Mauretania, northwest Africa, Lower Egypt, Israel, 
Jordan, Aden and Sokotra should be referred to R. h. sennaariense Fitzinger, 1866, 
a name thought for many years referable to R. microphyllum. Rhinopoma hard- 
wickei cy stops Thomas, 1903, to which all specimens from Egypt, the Sudan and 
Ethiopia were formerly referred is considered by Kock to be a smaller sub- 
species inhabiting Middle Egypt and extending westwards to Hoggar and Air in the 
Sahara : the small R. h. macinnesi Hayman, 1937 is thought by Kock to extend from 
northern Kenya and the southern Sudan to Somalia and to Eritrea in Ethiopia. 
These specimens from eastern Ethiopia are similar in size (length of forearm in 
thirteen examples 52-9-56-6 (55-0)) to those which this author refers to sennaariense 
from Khartoum and are considerably larger than macinnesi (length of forearm 
46-6-48-4, according to Kock (pp. 45, 50)). 



BATS FROM ETHIOPIA 31 

COLLECTION AND FIELD NOTES. The living bats were all shot or netted inside 
some small lava blister caves in the middle of the day. These provided the only 
shade and cool shelter in an otherwise hot, dry and very open terrain, covered with 
sparse thorn scrub, boulders and dry grass. Large numbers of Rhinopoma were 
found in these caves, often hanging from the roof in small solid masses of a dozen or 
more individuals. A sample of the bats was collected at random: the colonies 
contained both sexes, but no signs of any young were seen. Rhinopoma was also 
well represented among skeletal material removed from the floors of two of the 
lava caves. 



Taphozous perforatus haedinus Thomas 1915 

SPECIMENS, (i) Thirteen males, eight females. "Forward Base Two", 10 km 
west of Mabil, Blue Nile Gorge. 10 19' N, 36 45' E, alt. c. 1,300 m. 19 August 
1968. 

(2) One male, three females. North Bank of Awash River, Awash National 
Park, Shoa. 08 50' N, 40 01' E, alt. 1,000 m. 25-27 September 1968. 

TAXONOMIC NOTES. These specimens do not support the view (Harrison, 1961 : 
150; 1962 : 763) that two closely related species of the Taphozous perforatus group 
exist in Africa, one, perforatus, with dark wings, the other, Sudani, with pale or 
whitish wings, the two being separated also by a small difference in the size of the 
braincase, that of Sudani being on the whole very slightly larger than that of 
perforatus. As noted by Kock (1969 : 74), who also rejects this view, the pigmenta- 
tion of the wing membrane is variable. A number of the Ethiopian specimens have 
dusky wing membranes but in others the wing membranes are translucent, especially 
distally, and in this respect are closely similar to Sudani. Similarly, the dimensions 
of the braincase in the Ethiopian specimens bridge the narrow interval separating 
perforatus from Sudani. It is evident, therefore, that but a single species must be 
recognized, a course adopted by Rosevear (1965 : 151) and by Kock (1969 : 74) 
but counter to the opinion of Harrison (1961 : 150, 1962 : 763), who divided the 
group into two species, perforatus having a northern and eastern distribution in 
Africa, with Sudani distributed from the Sudan to southern Africa. 

Subspecific classification in T. perforatus is less clear, and final clarification must, 
it seems, await the advent of more material. The collections now available in the 
British Museum (Natural History) suggest that a pale subspecies (T. p. perforatus 
E. Geoffrey, 1818) occurring in northeastern Pakistan and in Egypt is replaced in 
the Sudan and Congo by a darker, slightly larger subspecies (T. p. Sudani Thomas, 
1915) which in turn in Ethiopia and Kenya gives way to an equally dark but slightly 
smaller subspecies (T. p. haedinus Thomas, 1915 : according to Kock (1969 : 80) 
Taphozous maritimus Heuglin, 1877 may be a prior name) extending to Somalia, 
Eritrea and southwestern Arabia. Specimens from southwestern Arabia are 
generally referred to T. p. haedinus but on the whole are a little paler than are 
those from Ethiopia and Kenya and thus tend towards T. p. perforatus: a small 
number of specimens reported as T. perforatus by Harrison (1968 : 323) from Oman 



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BATS FROM ETHIOPIA 33 

are said by that author to be distinctly paler than T. p. haedinus and hardly distin- 
guishable in colour from material from Sudan and Cutch. Specimens from Northern 
Nigeria have been separated as a distinct subspecies, T. p. swirae Harrison, 1962, 
on account of their greyish coloration : Kock (1969 : 80) lists this as a synonym of 
T. p. perforatus but thinks that there is a possibility that there may exist a distinct 
western subspecies, for which Taphozous senegalensis Desmarest, 1820 would be the 
earliest name, with swirae in synonymy. Extending in West Africa to Senegal, 
Ghana, Mali and the Cameroon, T. perforatus is known also from Tanzania, Botswana 
and Rhodesia, whence Harrison (1962 : 763) has described a small form, 
T. p. australis (preoccupied: as a subspecies of Sudani] subsequently renamed 
T. p. rhodesiae (Harrison, 19643. : 2) and considered a synonym of T. p. Sudani by 
Kock (1969 : 8l). Measurements of T. perforatus appear in Table i. 

COLLECTION AND FIELD NOTES. The specimens obtained near Forward Base Two 
all came from some small caves which lay at the head of a stream gulley, high up the 
side of the Nile Gorge. The gulley was steep, vertical in places, thickly over-grown 
and passed through dense thorn scrub covering the hillsides. The caves themselves 
were shallow and although dim light penetrated almost to their furthest extremities, 
they provided cool shelter from the sun. 

Large numbers of Taphozous were found together with single specimens of Nycteris 
thebaica and Hipposideros ruber. The cave floor was thickly carpeted with guano, 
and in one area, littered with the wings of large insects. These are presumed to repre- 
sent the remains of insect prey caught in flight by the bats and taken back to the 
roost where the thick bodies were eaten and the wings discarded. From their size and 
abundance it is assumed that these insects were the prey of Taphozous, rather than 
of the two smaller and less numerous bats. An endeavour was made to collect all 
wings in reasonable condition from the pile and these have been identified for the 
most part by Mr. Alan Brindle of the Manchester Museum, the remainder having 
been examined at the British Museum (Natural History) : the species encountered 
are listed in Table 2, with their relative abundance. 

An attempt was made to catch all the bats in the colony and in all, 40 Taphozous 
were obtained, leaving behind a few individuals that had retreated to inaccessible 
parts of the cave. All the bats were sexed and weighed in the field, then 21 of the 
Taphozous sample were released. The sex ratio in the full sample was 19 males: 

21 females. Weights ranged from 17 gms to 23 gms, with only two animals exceeding 

22 gms. The average weight was 19-5 gms with no significant sex difference. 
From the narrow size range, and the presence of both sexes in equal numbers and 
similar body weights it is evident that this was not a nursing colony and that 
breeding must occur at some other time of year, perhaps in a different place. 

The Taphozous collected in the Awash National Park were just a small sample 
taken to record their presence in the lava bubble caves. Here again the cavities 
were light throughout, but provided shelter and shade in an otherwise very open 
habitat. 



34 J- E. HILL & P. MORRIS 

TABLE 2 

Identities and abundance of insect wings from a roost of Taphozous perforatus 
Species identified Number obtained 

Orthoptera 

Tettigoniidae 

Diogena fausta (Burmeister) i 
Mantodea 

Tarachodes sp. i 

Polispilota aeruginosa (Goeze) i 
Lepidoptera 
Saturnidae 

Nudaurelea macrophthalma Ky 2 

Gyanisa maja Klug i 
Sphingidae 

Agrius convolvuli (Linnaeus) 35 

Hippotion eson (Cramer) 7 

Hippotion osiris (Dalman) i 

Hippotion celerio (Linnaeus) i 

Nephele peneus (Cramer) i 

Platysphinx stigmatica (Mabille) 2 

Euchloron megaera (Linnaeus) i 

Nycteris thebaica labiata (Heuglin, 1861) 

SPECIMENS, (i) One male, three females. Mabil, Blue Nile Gorge. 10 20' N, 
36 45' E, alt. c. 1,200 m. 18 August 1968. 

(2) Two females, one male. "Forward Base Two", 10 km west of Mabil, Blue 
Nile Gorge. 10 19' N, 36 45' E, alt. c. 1,000 m. 18-19 August 1968. 

TAXONOMIC NOTES. These specimens are similar in size (length of forearm 
41-45) to examples of N. t. thebaica from Egypt and from other localities (Gondar; 
Gallabat; Ghibbey Valley, o8i5'N., 3755'E.) in western Ethiopia but are 
slightly greyer ventrally. For this reason they are referred provisionally to 
N. t. labiata from Keren. The subspecies of N. thebiaca are discussed by Kock 
(1969 : 98). 

COLLECTION AND FIELD NOTES. The Mabil specimens were shot in the roof of a 
disused hut standing on a hilltop in open cultivated ground. One of the Forward 
Base Two specimens was taken from the same small caves as the large sample of 
Taphozous (above), the others from a similar little rock shelter lower down the 
same river gulley. 

ADDITIONAL SPECIMENS. The Sandhurst Ethiopian Expedition, 1966 also 
obtained N. thebaica at a locality northeast of Lake Chamo, southern Ethiopia, 



BATS FROM ETHIOPIA 35 

where specimens were found sleeping separately in deep, shady pits in the ground. 
Skulls of Nycteris sp. were also present among the cave floor bone debris collected 
from certain lava blister caves in the Awash National Park in September 1968. 

Cardioderma cor (Peters, 1872) 

SPECIMENS. Skeletal material. Awash National Park, Shoa. o85o'N, 40 
01 ' E, alt. c. 1,000 m. 28 September 1968. 

COLLECTION AND FIELD NOTES. Collected among large numbers of mammal 
bones found on the floor of the lava blister caves. 

ADDITIONAL SPECIMEN. A skin, thought to be Cardioderma, formed part of a 
small collection of local mammals held at the HQ of the Awash National Park. 

Rhinolophus clivosus acrotis Heuglin, 1861 

SPECIMENS. Skeletal material. Awash National Park, Shoa. o85o'N, 40 
01' E, alt. c. 1,000 m. 28 September 1968. 

COLLECTION AND FIELD NOTES. Subfossil cave remains, as for previous species. 

Rhinolophus landeri dobsoni Thomas, 1904 

SPECIMEN. One male. Temporary Base, mouth of Azir River, Blue Nile Gorge. 
10 29' N, 36 25' E. alt. 1,000 m. 20 August 1968. 

TAXONOMIC NOTES. This specimen agrees in size with the type specimen of 
dobsoni from Kordofan and with a small series from the Sudan and western Ethiopia 
in the collections of the British Museum (Natural History). The series as a whole 
confirms the measurements of Kock (1969 : 175) and shows that specimens from the 
Sudan and from Ethiopia are generally smaller (length of forearm in 12 examples 
41-5-44-6 (42-8) than those from Malawi (R. I. lobatus: length of forearm in 16 
examples 42-4-46-6 (44-8)), or from Tanzania (length of forearm in 5 examples 
44-4-46-0 (45-1)). Specimens from Zambia are slightly larger than those from 
Malawi (length of forearm in n examples 42-5-47-8 (45-5)) but from Kenya are 
smaller (length of forearm in 10 examples 41-1-45-8 (43-9)), approaching the 
northern subspecies in size. 

COLLECTION AND FIELD NOTES. Caught in hand-held mist net, 20.05 hrs on 
open river bank beside tall trees. 

Rhinolophus hipposideros minimus Heuglin, 1861 

SPECIMEN. One female. Lake Baa-sa-ka, Awash, Shoa. 08 50' N, 40 01' E, 
alt. c. 1,000 m. 26 September 1968. 

TAXONOMIC NOTES. This specimen appears to be the second of R. h. minimus to 
be recorded from Ethiopia, the subspecies being known in northeastern Africa from 
Keren (the type locality) and from Sennaar, Sudan (Andersen, 1904 : 455). The 



36 J. E. HILL & P. MORRIS 

specimen from Shoa agrees with the Sennaar example excepting only that it has 
narrower zygomata and has the anterior upper premolar (pm 2 ) smaller, more 
rounded and less angular in cross-section. Measurements (the Sennaar example 
in parentheses): length of forearm 367 (357); total length of skull to canine 14-4 
(14-5); condylocanine length 12-6 (127); rostral width 3-4 (3-3); zygomatic width 
6'7 (7 -5); least interorbital width 1-4 (1-5); width of braincase 6-2 (6-3); mastoid 
width 6-9 (7-1) ; c - m3 5-0 (5-0) ; c - m 3 5-3 (5-3). 

COLLECTION AND FIELD NOTES. Caught after dark (20.00 hrs) in a hand-held 
mist net as it flew low over the open, muddy lake shore. 



Rhinolophus simulator Anderson, 1904 

SPECIMEN. One female ; B.M. 64.854. Three miles south of Goba, Bale Province, 
(presumably about 5 30' N, 40 05' E P.M.) c. 3,000 m. 1962. 

TAXONOMIC NOTES. This specimen was first identified in 1964 on accession to the 
collections of the British Museum (Natural History) as an example of R. hipposideros 
minimus, but further examination in the course of reporting the Great Abbai 
material shows that although in the features of the noseleaf it clearly resembles 
this taxon, it is much too large, particularly cranially, to represent it. It is referred 
to the rather larger R. simulator, not hitherto reported from any more northerly 
locality than southern Tanzania, although Dr. K. Koopman (in litt.) of the American 
Museum of Natural History has identified specimens from western Kenya and the 
southern Sudan with this species. 

The Ethiopian specimen agrees closely with simulator in the structure of the sella, 
which is wide and very slightly constricted at a point a little above its centre and in 
its low, rounded connecting process which rises slightly above the rounded tip of the 
sella. The lateral margins of the lancet are slightly concave and the lancet itself is 
rounded towards the tip to f6rm a broad point. Apart from its generally smaller 
size, the skull agrees closely with that of simulator, with prominent rostral swellings, 
a shallow rostral sulcus, the anterior upper premolar (pm 2 ) in the toothrow and 
with the second lower premolar (pm 3 ) minute and extruded. Like simulator, it 
differs from R. swinnyi and R. denti (perhaps conspecific) in larger rostral swellings, 
the presence of a rostral sulcus and in having a longer supraorbital region. Rhino- 
lophus alticolus Sanborn, 1936 from the Cameroon is also very like simulator, differing 
chiefly in slightly larger size, rather less acuminate lancet and in having the mesop- 
terygoid fossa a little wider. There can be little doubt that simulator and alticolus 
are conspecific : the single specimen from Ethiopia points to the possible existence of 
a generally smaller montane subspecies in that region, a matter to be resolved by 
further specimens. The prior name is simulator by many years: alticolus, first 
described as a subspecies of R. alcyone, has evidently no close affinity with that 
species. Measurements of R. simulator are compared in Table 3. 

COLLECTION AND FIELD NOTES. Collected by P. M. Barrer and presented by the 
Imperial College (University of London) Ethiopian Expedition 1962. 



BATS FROM ETHIOPIA 



37 



TABLE 3 

Measurements of Rhinolophus simulator 



1 




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7-0 


68.1000 


$ 


44'5 


18-8 


16-4 


4'4 


9-0 


2-2 


8-6 


9-2 


6-5 


11-7 


6-9 


68.1001 


<J 


42-9 


18-3 


16-1 


4-6 


8-9 


2-2 


8-4 


9-1 


6-4 


"'5 


6-7 


68. 1 002 


6* 


4 2 -9 


18-5 


16-4 


4'5 


9-0 


2-2 


8-3 


9-0 


6-5 


n-3 


6-9 


14.6.13.2 


cJ 


43 ' 








4'3 














6-6 





7-0 


11.4.23.1 


- 


44-2 


18.0 


15-9 


4-4 





2-2 


8-4 


9-0 


6-2 





6-5 


Jf?. simulator 


























64.854 




42-6 


17-5 


15-4 


4'3 


8-5 


2-0 


7'7 


8-4 


6-0 





6-4 


R. s. alticolus 


























56.187 


6* 


46*3 


18-7 


16-7 


4-9 





2-2 





9-1 


6-8 





7'3 


56.188 


$ 


46*5 


18-6 


16-3 


4-8 


8-9 


2-3 


8-1 


9-0 


6-6 





7-0 


68.895 


5 


45'5 


























n-8 


7-2 


68.896 


6* 


45-8 


19-1 


16-9 


4'9 


9'3 


2-2 


8-3 


9-3 


6-9 


1 1 -9 


7-2 


68.897 


(J 


46-2 


18-8 


16-7 


4'9 


9'3 


2< 3 


8-3 


9-3 


6-7 


11-7 


7'3 


68.898 


c? 


44'9 


18-8 





4'9 


8-9 


2-3 








6-8 


11-7 


7-2 



Rhodesia 



Transvaal 
Zambia 



Malawi 
Tanzania 



Ethiopia 



Cameroon 



Rhinolophus fumigatus fumigatus Riippell, 1842 

SPECIMENS, (i) One female. Mouth of Fincha River, Blue Nile Gorge, 10 
3' N, 37 20' E, alt. 1,000 m. 12 August 1968. 

(2) One male. About 30 km southeast of "Portuguese Bridge", near Mota, 
Blue Nile Gorge, 11 20' N, 38 10' E, alt. c. 1,300 m. 22 September 1968. 

COLLECTION AND FIELD NOTES. Both specimens shot; the Fincha River one 
coming from a small rock fissure on a rocky hillside covered with thorn scrub and 
the other from a cave in a cliff beside the Nile. 



38 J. E. HILL & P. MORRIS 

Hipposideros caffer caffer (Sundevall, 1846) 

SPECIMEN. One male. "Forward Base Two", 10 km west of Mabil, Blue Nile 
Gorge, 10 19' N, 36 45' E, alt. c. 1,000 m. 18 August 1968. 

TAXONOMIC NOTES. Hitherto, many authors have followed Andersen (1906 : 275) 
in maintaining H. caffer as a polytypic species with a number of subspecies among 
which wide intergradation occurred. However, Hollister (1918 : 85) considered 
that two species, readily separable by size, were to be found together in East Africa, 
and, more recently, Lawrence (1964 : i) has suggested that the relative sizes of the 
narial compartments can be used as a specific distinction between these, the putative 
subspecies caffer and ruber. This author did not, however, attempt to allocate the 
various named forms hitherto ascribed to caffer beyond suggesting that centralis 
should be associated with ruber rather than with caffer. Kock (1969 : 130, 133; 
also discusses the classification of the group and concludes that two species can be 
recognized. Specimens from the Great Abbai Expedition and others from Ghana 
examined recently confirm the views of these authors and have prompted a further 
examination of the entire complex as it is represented in the collections of the 
British Museum (Natural History). The majority of specimens can be allocated 
readily to one or other of two groups, as Koopman (1966 : 158) has noted. One 
(caffer, angolensis, tephrus, Inanus) is composed of smaller (length of forearm usually 
less than 48), generally more greyish (in the dull phase) forms with small median 
posterior narial compartments and wide lateral inflations. Those allocated to the 
second group (ruber, centralis, guineensis, niapu] are generally larger (length of fore- 
arm usually greater than 48), browner (in the dull phase) and have larger median 
posterior narial compartments with narrow lateral inflations. There is evidently 
an ecological preference as is indicated by Verschuren (1957 : 354, 373 for centralis 
and nanus], Lawrence (1964 : 4) and Koopman (1966 : 158), the members of the first 
group occurring in the drier woodland and savannah regions, those of the second 
group in wetter, densely forested areas, as suggested by Brosset (1968 : 338). 
There exist, however, wide areas of sympatry on the fringes of the forest areas and 
this has led to the difficulties encountered when all of the named forms are considered 
to be subspecies of a single polytypic species (Hill, 1963 : 63). Members of either 
group may be readily recognized over most of Africa but in northern Angola and the 
Lower Congo the local representative (centralis} of the larger group is reduced in size 
and distinction from that (angolensis) of the smaller group is difficult. This circum- 
stance led Koopman (1966 : 158) to retain the concept of a single polytypic species. 
Both species occur in two colour phases : in the dull phase caffer is greyish and ruber 
brownish, while the bright phase of caffer is some shade of orange, of ruber more 
rufous. 

Hipposideros caffer is distributed throughout most of Africa excluding the central 
forested region from Morocco to Senegal, Sierra Leone, Ghana, Senegambia, Nigeria, 
northeastern Congo, Sudan, Ethiopia, Somalia, Kenya, Tanzania (including Zan- 
zibar), Pemba Island, Zambia, Rhodesia, Malawi, Natal, Transvaal, Cape Province, 
South West Africa, Angola and Gabon: outside Africa the species extends to the 
Yemen. The following subspecies may prove valid: 



BATS FROM ETHIOPIA 39 

Hipposideros coffer coffer (Sundevall, 1846) 

Mainly northeastern, eastern and southern Africa. 

Hipposideros coffer angolensis (Seabra, 1898) 

South West Africa; Angola; Gabon; Lower Congo. 

Hipposideros coffer tephrus Cabrera, 1906 

Northern and northwestern Africa ; drier regions of West Africa. 

Hipposideros coffer nanus J. A. Allen, 1917 
Northeastern Congo. 

It seems that aurantiacus de Beaux, 1924 from Somalia is based on an example of 
H. c. coffer in the red or brighter phase, while apparently braimo Monard, 1939 from 
Portuguese Guinea should be synonymized with H. c. tephrus (Aellen, 1956 : 26; 
Rosevear, 1965 : 226). Lawrence (1964 : 3) and Koopman (1965 : 10, 1966 : 158) 
agree that nanus J. A. Allen, 1917 is a subspecies of H. coffer. 

Specimens from the Sudan are referred to H. c. tephrus by Koopman (1965 : 10) 
and Kock (1969 : 130). The Ethiopian specimen from the Great Abbai collection, 
however, has a generally slightly larger skull than tephrus and consequently is 
referred to H. c. coffer. Measurements; length of forearm 46-9; greatest length of 
skull to canine 17-2; condylocanine length 14-9; rostral width 4-4; zygomatic 
width ; least interorbital width 2-8; width of braincase 8-6; mastoid width 9-4; 
c 1 -c 1 4-0; m 3 -m 3 5-8; c- m 3 5-8; length of mandible 10-1; c - m 3 6-4. 

COLLECTION AND FIELD NOTES. The single specimen was caught after dark 
(20.45 hrs) in a hand-held mist net as it flew low over a flat riverside sandbank in 
an area of thick bush. 

Hipposideros ruber centralis Andersen, 1906 
(Plate 3 (b)) 

SPECIMENS, (i) One female. Mouth of Fincha River, Blue Nile Gorge. 
10 03' N, 37 20' E, alt. 1,000 m. 12 August 1968. 

(2) Two males and two females. "Forward Base Two", 10 km west of Mabil, 
Blue Nile Gorge. 10 19' N, 36 45' E, alt. 1,000 m. 15-19 August 1968. 

(3) One male. "Forward Base Three", mouth of Didessa River, Blue Nile 
Gorge. 10 05' N, 35 38' E, alt. c. 1,000 m. 26 August 1968. 

TAXONOMIC NOTES. Reasons for regarding ruber as a distinct species rather than 
a subspecies of coffer are discussed above. It is of interest to note that specimens 
(B.M. 67.1135-1140) collected by the Sandhurst Ethiopian Expedition, 1966 at the 
southwest corner of Lake Abaya are also referable to H. r. centralis which clearly 
extends some distance into Ethiopia. 



4 o J. E. HILL & P. MORRIS 

Hipposideros ruber is distributed through the forests and savannahs of Ethiopia, 
the Sudan, Uganda, Kenya, Tanzania, Zambia, Angola, Gabon, Congo (Kinshasa), 
Congo (Brazzaville), Cameroon, Fernando Poo, Nigeria, Ghana, Gambia, Sierra 
Leone, Liberia, Spanish Guinea, Senegal, Sao Tome' Island and Principe Island. 
Possible subspecies are: 

Hipposideros ruber ruber (Noack, 1893) 

Tanzania, Zambia, Angola (Sanborn, 1950 : 58). 

Hipposideros ruber centralis Andersen, 1906 
Ethiopia, Sudan, Uganda, Kenya. 

Hipposideros ruber guineensis Andersen, 1906 
West Africa. 

Hipposideros ruber niapu J. A. Allen, 1917 
Northeastern Congo. 



Specimens reported by Aellen and Brosset (1968 : 447) as H. coffer from Congo 
(Brazzaville) seem likely to represent ruber (possibly H. r. centralis) : others reported 
from the Cameroon by Aellen (1952 : 72, 73) as H. c. caffer and H. c. angolensis 
seem from measurements to be referable to H. ruber. Aellen (loc. cit., pp. 74, 75) 
also records guineensis and ruber from the Cameroon, as subspecies of H. caffer. 

COLLECTION AND FIELD NOTES. The Fincha River specimen, and three of those 
from Forward Base Two were all caught after dark in a hand-held mist net as they 
flew low over riverside sand banks in thick scrub habitat. The specimen from the 
Didessa River was in rather less open habitat, flying around among riverside bushes 
at 21.05 hrs. The remaining H. ruber from Forward Base Two was netted in the 
same small caves as large numbers of Taphozous perforatus (q.v.). 

Two sharply contrasted colour phases are represented in the collection 
grey/brown and bright orange/red, with both forms being encountered at a single 
locality (Forward Base Two). 

Asellia tridens tridens (E. Geoffroy, 1818) 

SPECIMEN. Skeletal material. Awash National Park, Shoa. o85o'N, 40 
01' E, alt. c. 1,000 m. 28 September 1968. 

COLLECTION AND FIELD NOTES. Part of a collection of mammal bones removed 
from the floor of a lava blister cave. 



BATS FROM ETHIOPIA 41 

Asellia patrizii de Beaux, 1931 

SPECIMEN. One male. North bank of Awash River, Awash National Park, 
Shoa. 08 50' N, 40 01 ' E, alt. c. 1,000 m. 25 September 1968. 

TAXONOMIC NOTES. This species has been known hitherto only from Danakil, 
Ethiopia (the type locality) and from two other locations in Ethiopia, namely 
Assab, Eritrea and Entebebir Island, near Dahlak Kebir Island, off Massawa, 
Eritrea, the specimen recorded now from the Awash National Park being in fact the 
first to be received at the British Museum (Natural History). It demonstrates 
effectively the much smaller skull of patrizii when compared with tridens which has 
been obtained (vide supra) in cave remains from the same area. Harrison (1965 : 4) 
noted that specimens from Entedebir Island are slightly smaller than those 
recorded from the mainland and thought therefore that they might prove to be 
subspecifically separable. However, the specimen from the Awash National Park 
is very similar in size to those reported by Harrison and does not support this view. 
Measurements: length of forearm 40-1; greatest length of skull 14-7; condylobasal 
length 13-1; condylocanine length 12-7; zygomatic width 7-6; least interorbital 
width 1-9; width of braincase 6-1; mastoid width 7-1; c 1 - c 1 3-9; m 3 -m 3 5-2; 

COLLECTION AND FIELD NOTES. Shot hanging from the roof of a lava blister cave 
in very dry open, rocky terrain, 1500 hrs. 

Triaenops persicus afer Peters, 1877 
(Plate 3 (c)) 

SPECIMENS, (i) One male and one female. "Forward Base Three", mouth of 
Didessa River, Blue Nile Gorge. 10 05 'N, 35 38' E, alt. c. 1,000 m. 28 August 
1968. 

(2) One female. Awash National Park, Shoa. o85o'N, 4ooi'E, alt. c. 
1,000 m. 28 September 1968. 

TAXONOMIC NOTES. Although recorded from Somalia and Kenya, these speci- 
mens appear to be the first of Triaenops to be reported from Ethiopia. A rather 
larger subspecies, T. p. majusculus, has been described by Aellen and Brosset 
(1968 : 450) from the Congo (Brazzaville). 

COLLECTION AND FIELD NOTES. Both specimens obtained at Forward Base 
Three were caught with a hand-held mist net. The male was flying low over a 
maize plot on the bank of the Nile at 19.50 hrs, the female was flying low over the 
river itself at 22.00 hrs. The habitat is dense trees and bush with areas of maize 
cultivation, sharply contrasting with the dry, open Awash locality where the other 
specimen was shot in one of the lava blister caves in the middle of the day. 

ADDITIONAL SPECIMEN. A further specimen (B.M. 69.875) collected by Mr. C. 
Buer and presented by Dr. M. J. Largen has been examined recently. It was 
obtained on the main road between Lake Langano and Addis Ababa. 



42 J. E. HILL & P. MORRIS 

Pipistrellus kuhlii fuscatus Thomas, 1901 

SPECIMEN. One female. Ghimbi, Wollega. 09 10' N, 35 50' E, alt. 2,150 m. 
31 September 1968. 

TAXONOMIC NOTES. Mertens (1925 : 22) pointed out that africanus Ruppell, 
1842 from Shoa is very similar to f^lscatus Thomas, 1901 from Kenya, for which it 
is considered a prior name by Kock (1969 : 168). 

COLLECTION AND FIELD NOTES. This specimen was obtained "from a house" 
(presumably in the town of Ghimbi), no further information is available. 

Pipistrellus nanus (Peters, 1852) 

SPECIMENS, (i) One male and two females. Sabeta, Shoa. o855'N, 38 
40' E, alt. 2,500 m. August 1968. 

(2) One male. Ghimbi, Wollega. 09 10' N, 355o'E, alt. 2,150 m. 2 Sep- 
tember 1968. 

COLLECTION AND FIELD NOTES. The Sabeta specimens were collected by local 
children from the axils of banana leaves. The Ghimbi animal was found by a 
local boy, but no further details are available. 

Eptesicus somalicus somalicus (Thomas, 1901) 

SPECIMENS, (i) One male and one female. Mouth of Fincha River, Blue Nile 
Gorge. 10 03' N, 37 20' E, alt. c. 1,000 m. 12 August 1968. 

(2) One immature female. "Forward Base Two", 10 km west of Mabil, Blue 
Nile Gorge. 10 19' N, 36 45' E, alt. c. 1,000 m. 15 August 1968. 

COLLECTION AND FIELD NOTES. All specimens caught flying low over riverside 
sandbanks in thick bush habitat, between 20.00 hrs and 20.30 hrs using hand-held 
mist nets. 

ADDITIONAL SPECIMEN. A further specimen (B.M. 67.2164) collected northeast 
of Lake Abaya by the Sandhurst Ethiopian Expedition 1964 is referable to the 
rather larger species E. capensis. 

Glauconycteris variegata variegata (Tomes, 1861) 

SPECIMEN. One female, "Forward Base Two", 10 km west of Mabil, Blue Nile 
Gorge. 10 19' N, 36 45' E, alt. c. 1,000 m. 18 August 1968. 

TAXONOMIC NOTES. This specimen is the first of Glauconycteris to be recorded 
from Ethiopia. It has large, massive canines and cheek teeth which agree closely 
with G. v. variegata rather than with the Sudanese G. v. phalaena in which the den- 
tition is less massive. 



BATS FROM ETHIOPIA 



43 



COLLECTION AND FIELD NOTES. By chance this specimen was found dead before 
being eaten by scavengers. It was lying at the river's edge among boulders which 
are subject to frequent inundation. The surrounding habitat consists of tall trees 
and dense scrub. % 

The following new species is described below by the senior author: 

Myotis morrisi Hill, sp. nov. 
(Plates i, 2 (a, b)) 

SPECIMEN. HOLOTYPE. B.M. 70.488. Adult female. A flat, card-mounted skin 
with skull; collector's number Aio7. "Forward Base Three", mouth of Didessa 
River, Blue Nile Gorge. 10 05' N, 35 38' E, alt. c. 1,000 m. 28 August 1968. 

OTHER MATERIAL: none. 

TAXONOMIC NOTES. DIAGNOSIS. Similar to Myotis tricolor (Temminck, 1832) 
of eastern Africa but differing from this species in its generally more orange dorsal 
coloration; unicolored and not bicolored ventral pelage; elongate, narrower skull 
with uninflated braincase and supraorbital region, the braincase shorter, more 
globular and markedly narrower than in M. tricolor, the rostrum proportionately 
longer and less broadened than in that species. Dentition less massive than in 
M. tricolor ; second upper premolar (pm 3 ) relatively larger and incompletely intruded 
from the toothrow; second lower premolar (pms) relatively larger, not compressed 
between pm 2 and pnu; posterior upper premolar (pm 4 ) with narrower lingual shelf, 
separated from the lingual shelf of m 1 by a wider interspace. 

DESCRIPTION. Of moderate size (length of forearm 45-4) for the genus; anterior 
margin of ear smoothly convex, posterior margin concavely emarginated in its distal 
half, the proximal half convex. Tragus long, its length equal to one half of the 
length of the ear, tapered, with slender, rounded tip directed slightly posteriorly; 
anterior margin faintly convex, especially distally, posterior margin concave distally, 
convex proximally, slightly serrated, a small rounded lobe at base beneath an acute, 
angular emargination just below widest point of tragus. Wing inserted at base of 
first toe ; calcar strongly developed, extending along almost one half of the posterior 
margin of the tail membrane ; no obvious post-calcareal lobe. 

Pelage woolly, dorsally overall orange brown, individual hairs tricolored, the basal 
quarter blackish brown, most of remainder creamy white, hairs tipped terminally 
with bright orange brown. Dorsal pelage extending narrowly on to the wing 
membrane and on to the tail membrane for nearly one half of its width. Ventral 
pelage unicolorous dull creamy white, tinged faintly with brown on chin and flanks. 
Wing membrane generally black but antebrachium and endopatagium pale orange 
yellow, a narrow band of the same colour extending across the membrane immedi- 
ately behind the forearm; anterior edge of membrane and area between first and 
second metacarpals and tail membrane similarly coloured. Tibia flanked by a 
narrow band of orange hairs on wing membrane and a wider band of similar hairs on 
tail membrane. Toes with a sparse covering of long orange hairs. 



44 J- E. HILL & P. MORRIS 

Skull (Plates i, 2 (a)) elongate, the braincase not especially inflated; rostrum 
narrow, the supraorbital region not expanded, the supraorbital ridge forming an 
uninterrupted curve; a shallow, narrow median rostral depression. Narial emar- 
gination narrow, V-shaped posteriorly, its apex rounded. Palate long, narrow, 
anterior palatal emargination rounded posteriorly, extending almost to a line 
joining the centres of the canines; narrow post-palatal extension; shallow basi- 
occipital pits. Inner upper incisor (i 2 ) longer than wide, bicuspid, with strong 
angular anterior cusp ; smaller posterior cusp extending for two thirds of the height 
of the anterior cusp. Outer upper incisor (i 3 ) wider than long, bicuspid, closely 
appressed to inner tooth, cusps lying transversely to line of toothrow. Outer cusp 
the larger, rising from a narrow cingulum shelf to a height equal to that of the 
posterior cusp of inner tooth ; supported internally by a smaller secondary cusp for 
two thirds of its height, the tooth somewhat hollowed internally, separated from 
the canine by a short diastema. Anterior upper premolar (pm 2 ) about as high as i 3 , 
a little larger at base, in contact with canine, a slender pointed cusp with wide, 
strong cingulum. Second upper premolar (pm 3 ) rising slightly above cingulum of 
pm 2 , slightly less than one half its basal area, visible externally, a simple cusp rising 
from a narrow cingulum, slightly intruded from row but separating pm 2 and the 
posterior upper premolar (pm 4 ), in contact with these teeth. Posterior upper 
premolar (pm 4 ) with strong cusp and short, narrow lingual shelf with a wide inter- 
space, separating it from the lingual shelf of the first upper molar (m 1 ). Lower 
incisors imbricated, first (ii) and second (i 2 ) with four cusps, outer cusp of ii incipient, 
of 12 well developed but lower than inner cusps. Lower premolars not especially 
compressed: second (pm 3 ) rather more than one half the height of anterior tooth 
(pm 2 ) and approximately one half its basal area, in contact with pm 2 and posterior 
lower premolar (pm4), but not compressed or displaced, its length and width equal. 
The measurements of the new species are compared with those of M. tricolor in 
Table 4. 

REMARKS. In some respects M. morrisi resembles M. bocagei but is larger, has 
unicolored and not bicolored ventral pelage, a relatively longer rostrum lacking 
supraorbital inflation, narrower narial and anterior palatal emarginations and 
relatively narrower palate. Dentally the two species are closely similar but the 
teeth of M. morrisi are more generally massive than are those of M. bocagei and 
M. morrisi lacks a protoconule on the anterior ridge of the first and the second upper 
molars (described in M. bocagei by Harrison (1964^ : 135).) 

Tate (1941 : 539) reviewed the subgenera of Myotis and (p. 552) referred tricolor 
to the subgenus Selysius. However, both tricolor and morrisi approach the sub- 
genus Chrysopteron in dichromatic wing pattern, although this is less evident in 
tricolor, and in the presence of four lobes on the inner (ii) and second (i 2 ) lower 
incisors, but retain a relatively high braincase and concave frontal profile. I have 
much pleasure in associating with this new species the name of my co-author, 
Dr. Pat Morris, of Royal Holloway College, University of London, in recognition 
of his many services to the study of the Ethiopian fauna while with the Great Abbai 
Expedition. 



BATS FROM ETHIOPIA 



45 



Cu 
o 

3 

W 



II ! 

x jif <z 



jo 



* o 
W S 

' 



jo 



o ooooooooooooooo 



SSOJ013 



jo 



jo 



Of 



^o CH- Of Of *o *O Of Of Of Of 





:aw 



^ ^ 



O O O 



M IOIOIOIO 



46 J. E. HILL & P. MORRIS 

COLLECTION AND FIELD NOTES. The specimen was caught by Messrs. Yalden, 
Largen and King of the Great Abbai Expedition, using a hand-held mist net. The 
bat was flying after dark over the river Nile, near its north bank, at a height of 
about one metre. The surrounding habitat comprises mainly maize plots and thick 
bush. The specimen was photographed whilst still alive (Plate 2 (b)). 

Scotophilus sp. 

SPECIMENS, (i) One subadult male. Mouth of Fincha River, Blue Nile Gorge, 
10 03' N, 37 20' E, alt. 1,000 m. n August 1968. 

(2) One subadult male. "Forward Base Two", 10 km west of Mabil, Blue Nile 
Gorge, 10 19' N, 36 45' E, alt. c. 1,000 m. 15 August 1968. 

TAXONOMIC NOTES. No attempt has been made to allocate these subadult 
specimens to any of the named forms of Scotophilus in northeastern Africa. 

COLLECTION AND FIELD NOTES. Both individuals were found flying over riverside 
sandbanks at a height of about 2 metres, the surrounding habitat consisting mainly 
of dense scrub. The Fincha River specimen was shot in flight (with a pistol!) at 
19.15 hrs, the other animal was the only bat caught in a fixed, stationary mist net 
(19.30 hrs) on the whole Expedition, except for bats caught at their roosts. 

Miniopterus inflatus africanus Sanborn, 1936 

SPECIMEN. One female. North bank of Awash River, Awash National Park, 
Shoa. 08 50' N, 40 01 ' E, alt. c. 1,000 m. 23 September 1968. 

COLLECTION AND FIELD NOTES. The bat was flying fairly high (6-8 metres) 
among widely spaced acacia trees in open thorn scrub and grassland. It was shot 
in flight at 23.00 hrs. 

Otomops martiensseni martiensseni (Matschie, 1897) 

SPECIMEN. One female, B.M. 69.1256. French Somaliland (Territory Afars and 
Issas). 11 46' N, 42 39' E, alt. 1,471 m. 8 August 1967. 

TAXONOMIC NOTES. This specimen, collected by the Sandhurst French Somali- 
land Expedition, records 0. martiensseni for the first time from French Somaliland 
and represents a wide extension of range from Kenya. 

Tadarida pumila (Cretzschmar, 1830) 

SPECIMENS, (i) One male and eight females (three of them immature) . North 
bank of Awash river, Awash National Park, Shoa. o85o'N, 4ooi'E, alt. c. 
1,000 m. 25-26 September 1968. 

(2) Two males and four females. Ghimbi, Wollega. 09 10' N, 35 50' E, alt. 
2,150 m. 1-4 September 1968. 



BATS FROM ETHIOPIA 



47 



COLLECTION AND FIELD NOTES. The Awash specimens formed part of a roost of 
bats, living behind the loose bark of a dead tree. A sample of the animals was 
collected in the late afternoon using a mist net fixed to the tree trunk. Four of the 
Ghimbi animals (two females and two males) were living in the eaves and under the 
corrugated iron sheeting of the Mission Church roof, one of the others is recorded as 
coming from a house and the exact origin of the sixth Ghimbi specimen is not known, 
though it is likely to have been collected with the rest. 

These two localities for T. pumila (Awash and Ghimbi) could not be more contrast- 
ing and suggest a lack of strict habitat requirements for this bat. The Awash 
locality was very hot, dry, open bush whereas Ghimbi is 1,200 m higher on the cool, 
wet, heavily cultivated highland plateau. The only similarities between the two 
roosts are that the bats were living close together in a narrow, cramped space and 
both sexes were present in each colony. 



Tadarida nigeriae nigeriae (Thomas, 1913) 
(Plate 3 (d)) 

SPECIMEN. One male. North bank of Awash River, Awash National Park, 
Shoa. 08 50' N, 40 01' E, alt. c. 1,000 m. 26 September 1968. 

TAXONOMIC NOTES. There is but one previous record of T. nigeriae from Ethiopia 
(Ingersol, 1968 : 60), from the Gota River in the eastern part of the country. The 
species has been known hitherto from no locations nearer to Ethiopia than the 
northeastern Congo and southeastern Tanzania. 

COLLECTION AND FIELD NOTES. This single specimen was among a sample taken 
from the roost of T. pumila mentioned above. It seems peculiar that two species 
should be living behind the same small piece of tree bark in a mixed colony. 



Tadarida africana (Dobson, 1876) 

SPECIMEN, (i) One male B.M. 28.1.11.40. Fatam river, Great Abbai (=Blue 
Nile), c. 70 km south of Lake Tana, approx. 10 25' N, 37 oo' E. alt. c. 1,900 m. 
17 March 1927. 

(2) One specimen B.M. 69.884. ? Vicinity of Addis Ababa. 

TAXONOMIC NOTES. These specimens are the first of this large molossid to be 
reported from Ethiopia, reported hitherto from no nearer locality than southwestern 
Kenya. 

COLLECTION AND FIELD NOTES. The Fatam River specimen was collected by 
Major R. E. Cheesman during one of his survey visits to the Nile Gorge. The other 
individual was found dead on a telegraph wire by employees of the Imperial High- 
way Authority and presented by Dr. M. Largen of the Haile Selassie University, 
Addis Ababa. 



48 J. E. HILL & P. MORRIS 

Tadarida acetabulosus natalensis (A. Smith, 1847) 

SPECIMEN. One female B.M. 6.11.1.9. Given on the specimen label as "between 
Shoa and Lake Rudolf" Southern Ethiopia. 

TAXONOMIC NOTES. This single specimen represents a wide extension of range 
for T. acetabulosus, known hitherto from Madagascar, Mauritius, Reunion and Natal, 
the latter record by A. Smith being hitherto the only evidence of the occurrence of 
the species on the African mainland. Although Thomas identified the specimen 
correctly upon its arrival at the British Museum (Natural History) in 1906 the 
record has remained unpublished and the specimen undisturbed in the collections 
until it was noted by Mr. R. W. Hayman in 1965. 

SUMMARY 

The Great Abbai Expedition obtained 115 specimens of bats (including cave 
remains), chiefly from the Blue Nile Gorge or from the Awash National Park. One 
specimen from the Blue Nile Gorge proves to represent a new species closely allied to 
Myotis tricolor, for which the name Myotis morrisi is proposed. Specimens in the 
collections of the British Museum (Natural History) and also others collected by the 
Expedition confirm the presence of Micropteropus pusillus in Ethiopia; Triaenops 
persicus afer and Glauconycteris variegata variegata were obtained for the first time 
in Ethiopia by the Great Abbai Expedition, which also obtained further specimens 
of Asellia patrizii and Talarida nigeriae nigeriae; a few specimens in the British 
Museum (Natural History) obtained from other sources and reported in this paper 
furnish the first Ethiopian records of Rhinolophus simulator, Tadarida africana and 
Tadarida acetabulosus, and of Otomops martiensseni in French Somaliland (Territory 
Afars and Issas). The classification of Taphozous perforatus, Hipposideros caffer 
and Hipposideros ruber is reviewed. Rhinolophus simulator Andersen, 1904 and 
Rhinolophus alticolus Sanborn, 1936 are considered to be conspecific. 

REFERENCES 

AELLEN, V. 1952. Contribution a l'6tude des chiropteres du Cameroun. Mem. Soc. neuchat. 
Sci. nat. 8: 1-121, 26 figs., map. 

1956. Le Pare National du Niokolo-Koba (premier fascicule). II. Chiropteres. 

Mem. Inst. fr. Afr. noire No. 48 : 25-34, 5 tabs. 

& BROSSET, A. 1968. Chiropteres du Sud du Congo (Brazzaville). Revue suisse Zool. 

75 : 435-458, 2 figs., i pi. 

ANDERSEN, K. 1904. On von Heuglin's, Riippells's and Sundevall's types of African Rhino- 
lophi. Ann. Mag. nat. Hist. (7), 14 : 451-458. 

1906. On Hipposideros caffer Sund., and its closest allies : with some notes on H. fuli- 

ginosus, Temm. Ann. Mag. nat. Hist. (7) 17 : 269-282, i tab. 

1912. Catalogue of the Chiroptera in the collection of the British Museum. 2nd. ed. i. 

Megachiroptera. London. 

BLASHFORD-SNELL, J. N. 1970. Conquest of the Blue Nile. Georgl J. 136 : 42-60. 
BROSSET, A. 1968. La permutation du cycle saisonnier chez le chiroptere Hipposideros 
caffer, au voisinage de 1'Equateur. Biologia Gabon 4 : 325-341, 4 figs. 



BATS FROM ETHIOPIA 



49 



ELLERMAN, J. R. ( MORRISON-SCOTT, T. C. S. and HAYMAN, R. W. 1953. Southern African 

Mammals 1758 to 1951 : a reclassification. London. 
HARRISON, D. L. 1958. A new race of tomb bat, Taphozous perforatus E. Geoffrey, 1818, 

from northern Nigeria, with some observations on its breeding biology. Durban Mus. 

Novit. 5 : 143-149, i fig. 

1961. Notes on Southern and East African bats. Durban Mus. Novit. 6 : 149-152, i fig. 

- 1962. On bats collected on the Limpopo River, with the description of a new race of 
tomb bat, Taphozous sudani Thomas, 1915. Occ. Pap. natn. Mus. Sth. Rhod. No. 26 B : 
755-7 6 7. 3 n gs., 4 pis., 6 tabs. 

- i964a. Notes on some Southern Rhodesian Microchiroptera. Arnoldia, Bulawayo, 1; 

3 : i-3- 
- I9&4b. The mammals of Arabia. I. Insectivora; Chiroptera; Primates. London. 

- 1965. Remarks on some trident leaf -nosed bats (genus Asellia Gray, 1838) obtained by 
the Israel south Red Sea Expedition, 1962. Bull. Sea Fish. Res. Stn. Israel 38 : 23-5. 

- 1968. On three mammals new to the fauna of Oman, Arabia, with the description of a 
new subspecies of bat. Mammalia, 32 : 317-325, i pi., 3 tabs. 

HILL, J. E. 1963. A revision of the genus Hipposideros. Bull. Br. Mus. nat. Hist. (Zool.) 

11 : 1-129, 4 1 figs., 2 tabs. 
HOLLISTER, N. 1918. East African mammals in the United States National Museum. 

Part I. Insectivora, Chiroptera and Carnivora. Bull. U.S. natn. Mus. 99 : 1-194, 55 P^ s - 
INGERSOL, -R. H. 1968. The ecological stratification of mammals in the eastern Chercher 

Highlands of Harar Province, Ethiopia. Ph. D. Thesis, Oklahoma State University, i-viii, 

1-169, 28 figs., 12 tabs. 
KOCK, D. 1969. Die Fledermaus - Fauna des Sudan. Abh. senckenberg. naturforsch. Ges. 

521 : 1-238, 20 figs., 43 tabs. 
KOOPMAN, K. F. 1965. Status of forms described or recorded by J. A. Allen in "The American 

Museum Congo Expedition Collection of Bats." Am. Mus. Novit. No. 2219 : 1-34. 

1966. Taxonomic and distributional notes on Southern African bats. Puku 4 : 155-165. 
LAWRENCE B., 1964 Notes on the horshoe bats Hipposideros caffer, ruber and beatus 

Breviora, No. 207 : 1-5. 
MERTENS, R. 1925. Verzeichnis der Saugetier Typen des Senckenbergischen Museums. 

Senckenbergiana 7 : 18-37. 

ROSEVEAR, D. R. 1965. The bats of West Africa. London. 
SANBORN, C. C. 1950. Chiroptera from Dundo, Lunda, northeastern Angola. Publicoes 

cult. Co. Diam. Angola 10 : 51-62, 5 figs. 
TATE, G. H. H. 1941. Review of Myotis of Eurasia. Results of the Archbold Expeditions 

No. 39. Bull. Am. Mus. nat. Hist. 78 : 537-565, 2 figs. 
VERSCHUREN, J. 1957. Ecologie, biologic, et systematique des Chiropteres. Explor. Pare. 

Nat. Garamba, Miss, de Saeger. Inst. Pares. Nat. Congo Beige, Bruxelles, No. 7 : 1-473, 

173 figs., i pi., map. 



J. E. HILL, 

Department of Zoology, 

BRITISH MUSEUM (NATURAL HISTORY), 

CROMWELL ROAD, 

LONDON S.W.7. 

P. MORRIS, Ph.D., 

Department of Zoology, 

ROYAL HOLLOWAY COLLEGE (LTNIVERSITY OF LONDON), 

ENGLEFIELD GREEN, SURREY. 



PLATE i 
Myotis morrisi. Skull and mandible xy-5 



Bull. Br. Mus. nat. Hist. (Zool.) 21, a 



PLATE i 





PLATE 2 



(a) Myotis morrisi. Skull and mandible X7'5 

(b) Myotis morrisi. 



Bull. Br. Mus. nat. Hist. (Zool.) 21,2 



PLATE 2 





PLATE 



(a) Rhinopoma hardwickei sennaariense 

(b) Hipposideros ruber centralis 

(c) Triaenops persicus afer 

(d) Tadarida nigeriae nigeriae 



Bull. Br. Mus. nat. Hist. (Zool.) 21, 2 



PLATE 3 





(b) 



(d) 





Printed in England by Staples Printers Limited at their Kettering, Northants, establishment 










SOME SPECIES OF PARASITIC 
WORMS IN THE 'DISCOVERY' 
COLLECTIONS OBTAINED IN 
THE YEARS 1925-1936 



S. MARKOWSKI 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 21 No. 3 

LONDON: 1971 



ON SOME SPECIES OF PARASITIC WORMS 
IN THE 'DISCOVERY' COLLECTIONS 
OBTAINED IN THE YEARS 1925-1936 




BY 

STANISLAW MARKOWSKI 



Pp. 51-65; 20 Text-figures 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 21 No. 3 

LONDON: 1971 



THE BULLETIN OF THE BRITISH MUSEUM 

(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 21, No. 3 of the Zoological 
series. The abbreviated titles of periodicals cited follow 
those of the World List of Scientific Periodicals 



World List abbreviation 
Bull. Br. Mus. nat. Hist. (Zool.). 



Trustees of the British Museum (Natural History), 1971 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 10 September, 1971 Price 6op 



ON SOME SPECIES OF PARASITIC WORMS 
IN THE 'DISCOVERY' COLLECTIONS 
OBTAINED IN THE YEARS 1925-1936 

By S. MARKOWSKIf 



MATERIAL AND METHODS 

THE collection described below was made by 'Discovery' expeditions between the 
years 1925 and 1936 and consists of twenty-three samples of parasitic worms taken 
from the muscles and mesenteries, but mostly from the intestinal tract, of four 
species of fish (Euthynnus pelamis, Coryphaena sp., Chaenocephalus aceratus and 
Notothenia rossi), two species of birds (Phoebetria fusca and Chloephaga picta 
leucoptera), two species of seals (Leptonychotes weddelli, Hydrurga leptonyx) and a 
blue whale (Balaenoptera musculus). The material from seals was collected from 
six specimens of Leptonychotes weddelli and ten of Hydrurga leptonyx. 

In the material studied, twelve separate species of parasitic worms have been 
recognized. Of these, two forms of Cestodes seem to represent new species. In 
three cases, because of the juvenile condition of the specimens, generic determination 
only was possible. 

The bulk of the material was preserved in 4% formalin. Some specimens were 
prepared as whole mounts, having been stained with Mayer's paracarmine. Serial 
sections, cut at 8 (xm thick, were stained with Ehrlich's haematoxylin and counter 
stained with erythrosin. The hosts were taken at localities in the Southern Ocean, 
the precise localities are given with details of each particular species of parasite. 

The author takes this opportunity in expressing his thanks to the National 
Institute of Oceanography for entrusting this material to him and for providing 
the necessary scientific apparatus. 

Thanks are also due to Dr J. P. Harding, Keeper of the Zoology Department 
of the British Museum (Natural History) for providing the writer with accommodation 
and the loan of microscopical equipment, and to Mr S. Prudhoe, Mr J. W. Coles 
and Mr R. A. Bray of the same Museum for their assistance in the course of this 
investigation, as well as to Professor J. G. Baer, who kindly lent the original material 
of Hymenolepis bisaccata Fuhrmann, 1906 for comparison. 

fDr. Markowski died January 5, 1971 



54 



S. MARKOWSKI 




m 



42 



Sies of 
Nematodes 



peci 



O w 






t/5 rpl 

.s| 




S 


g-g 





* 8 






>SJ 


c^O 


i 


^^ 




"s E 


:S 




1 

^ "^ 


1.1 

"** ^ 




r^ <i, 

h 0?" 


o g 

oq-^ 









lashleyi Contracaecum 
'rfoliatus osculatum 


quadratum Contracaecum 
osculatum 




^ 




Is &, 


I 




o 


"> 


s 


s 


Tetrabothriiis 
heteroclitus 


Hymenolepis 
prudhoei sp. n 


Diplogonopont 
balaenopterae 


Diphyllobothri 
Glandicephalu 


Diphyllobothri 








2 










1 





O 173 
cfl O 
V 
'o 
O & 

^ i 







S 
8 

11 

S v 

^ ^ 



Notothen 
rossi 




^ 

C 

"S 

O M 
<^i vS 

o ^ 
* s 
A. < 



^ 
^1 

il 

IT 

J -S 

>4 

O 



-S, OQ 



alaenopt 
usculus 






9. 



yd 



PARASITIC WORMS IN THE 'DISCOVERY' COLLECTIONS 
ABBREVIATIONS USED IN THE FIGURES 



55 



c cuticula 

c.s. cirrus-sac 

ex excretory vessel 

l.m. longitudinal muscles 

m.ex. median excretory vessel 

o ovary 

s.m. subcuticular muscles 



t testis 

u uterus 

v vagina 

v.d. vas deferens 

v.g. vitelline glands 

v.s. vesicula seminalis 



SYSTEMATIC NOTES 



The material examined contains twelve species of parasitic worms belonging to 
four different groups and these are enumerated below. 



I. TREMATODA 

NOTOCOTYLIDAE Liihe, 1909 

i. Ogmogaster antarctica Johnston, 1931 

Host: Leptonychotes weddetti, intestine. Locality: Falkland Islands 15.7.1928. 

Several specimens of this trematode were found in the intestine of two Weddell 
seals. In one case they were attached to the walls of the intestine close to strobilae 
of Glandicephalus perfoliatus. 

Descriptions of this trematode have been given by Johnston (1931 and 1937). 



II. CESTODA 
TENTACULARIIDAE Poche, 1893 

2. Tentacularia (larvae) 

Host: Euthynnus pelamis: cysts in the abdominal muscles. Locality: i5io'N; 
i83o'W; 15.10.1925. 

Some six specimens are here recorded. Similar larval stages of a tetrarhynch 
were described from the same host-species by Rennie and Reid (1912). 



DIPHYLLOBOTHRIIDAE Liihe, 1910 
3. Diphyllobothrium lashleyi (Leiper and Atkinson, 1914) 

Host : Leptonychotes weddetti intestine. Localities : Falkland Island, and Grytviken, 
South Georgia; 15-17.7.1928. 

Great numbers of specimens were obtained from each of three seals. 



56 S. MARKOWSKI 

4. Diphyllobothrium quadratum (v. Linstow, 1892) 

Host: Hydmrga leptonyx, small intestine and rectum. Localities: Grytviken, 
South Sandwich Islands; South Orkney; 15.9, 18-22.1.1928 : 16.2.1931. The 
material was collected from ten seal-hosts, each showing a very heavy infestation. 



5. Diplogonoporus balaenopterae Loennberg, 1892 

Host: Balaenoptera musculus, intestine. Locality: 6i53'S, 8732'E, 27.1.1936 
('Southern Empress'). Large portions of strobila and few smaller fragments were 
found in this sample. 



6. Glandicephalus perfoliatus (Railliet and Henry, 1912) 

Host: Leptonychotes weddelli, intestine. Localities: Falkland Islands, 15.7.1928; 
Palmer Archipelago, 8.1.1935. 

Parts of the duodenum of two seal-hosts were found infested with this species. 
Detailed descriptions of the above-mentioned diphyllobothriid cestodes have been 
given in earlier papers (Markowski, 1952 and 1955). 



PTYCHOBOTHRIIDAE Liihe, 1902 
7. Bothriocephalusjanickiisp.nov. Figs 1-5 

Host: Coryphaena sp. ; stomach. Locality 2405'S, I546'N. 27.11.1925. 

Some twenty-one fragments and eight complete worms were examined. The 
strobila is about 8 cm long and very slender, about i mm broad. The scolex is 
very large in relation to the rest of the body, being 5 mm long and i mm broad. It 
is provided with a pair of groove-like bothridia. A neck was not observed (Fig. i). 

The excretory system consists of two pairs of longitudinal vessels, two individual 
canals at either side of the body. Of each pair the outward or dorsal vessel is 
about 5x5 (Am in diameter and the inward or ventral vessel about 
22-27 M- 111 x 14-21 (xm. Another single median longitudinal canal of about 
11-13 x 5-10 [/.m is situated at the right side of the cirrus-sac and the uterine 
opening. Its walls are thick and provided with cells arranged radially, as seen 
in the Figs 2 and 3. 

The longitudinal musculature consists of a very thin layer of fibres lying im- 
mediately beneath the body-cuticula and two well-developed layers of fibres in- 
serted in the parenchyma (Fig. 4). The cuticula is about 3-5 [zm thick. 

The sexually-mature segment is from 0-87 mm to 0-9 mm broad, as measured 
in transverse section. The genital pore, situated dorsally, leads into a shallow 



PARASITIC WORMS IN THE 'DISCOVERY' COLLECTIONS 



57 



genital atrium. The cirrus-sac, measured in the same place, is 160 [Am high and 
84 (Am broad, elongate, pyriform and situated dorsally (Fig. 2). There appear to 
be about 75 testes, but a more precise number, it has not been possible to determine. 
They are arranged in a single layer in the central part of the segment and measure 
about 40-50 x 30-35 [Am (Fig. 5). 

The ventrally-situated tocostoma or uterine pore leads into an atrium, which as 
measured in transverse section is 68 [Am high and 40 [Am wide (Fig. 3). 

The vitelline glands are disposed ventrally in a single layer between the longi- 






FIG. i. Bothriocephalus janickii sp. nov. : scolex. 

FIG. 2. Bothriocephalus janickii sp. nov.: cross-section of the segment showing cirrus-sac. 
FIG. 3. Bothriocephalus janickii sp. nov. : cross-section of the segment showing tocostoma. 



58 S. MARKOWSKI 

tudinal muscles, sometimes slightly overlapping into the lateral fields of the seg- 
ment. They are about 25-30 x 20 (xm (Fig. 4). The ovary is a deeply bilobed 
structure situated in the middle region of the segment. The eggs are 40-42 x 28- 
32 [xm. 

There are five species of Bothriocephalus occurring in fish-hosts in the Southern 
Hemisphere. However, the descriptions of some of them are very inadequate 
(Prudhoe, 1969). 

Bothriocephalus janickii sp. nov. differs from others quoted by Prudhoe (1969) 
with its unusually large scolex and extremely slender strobila. 

The species is named after the well-known Polish zoologist, the late Professor 
C. Janicki. 



HYMENOLEPIDIDAE Railliet & Henry, 1909 
8. Hymenolepis prudhoei sp. nov. Figs 6-14 

Host: Chloephaga picta leucoptera, rectum. Locality: Teal Inlet East Falkland, 

5-3-I927- 

Some sixty-five adult worms were collected. The length of the strobila in these 
specimens is about 9 cm and the width 4 mm. The scolex is about 125 ^m in 
length and 137 (xm in width (Fig. 6). The rostellar sac is unusually long in compari- 
son with the scolex, being about 187 (jun in length and 37-5 ^m in width. 

The rostellum bears a crown of eight hooks, each measuring 32 (xm. They have 








FIG. 4. Bothriocephalus janickii sp. nov.: cross-section of the segment showing the 

arrangement of the longitudinal muscles and vitellaria. 
FIG. 5. Bothriocephalus janickii sp. nov.: horizontal section of the segment. 



PARASITIC WORMS IN THE 'DISCOVERY' COLLECTIONS 



59 



a long well-developed blade, and a very thick, club-shaped handle. The guard is 
weakly developed (Fig. 7). A neck was not observed. The segments are short, 
elongate transversely. 

The three testes, 270-212 X 112-5-125 (xm are situated posteriorly across the 
segment in a single row (Figs 8 and 14). The vesicula seminalis is large and situated 
in the anterior region of the segment (Fig. 9). It opens with a coiled duct into the 
cirrus-sac, which measures about 130 [jan in length and 85 (j.m in width, and is 
provided with thick muscular walls (Fig. 10). 

The cirrus is armed with a smooth stylet, which may be observed protruding from 
the genital opening. 

The ovary is more or less rounded and connected with the ramifying uterus, 
which in the gravid segments occupies the whole proglottis. The embryo is en- 
closed in two membranes. The size of the outer membrane of the egg is 45 x 40 [j.m, 
the inner membrane is 37 x 28 [zm and the embryo itself 20 x 28 (xm (Fig. 13). 
Embryonic hooks were not observed. 

The longitudinal muscles are well developed and form two layers, of these the 
outer one is more strongly developed (Fig. 12). 





I. 8 



^'^{^J^^^^^^^A-^: >: J' 



FIG. 6. Hymenolepis prudhoei sp. nov. : scolex. 
FIG. 7. Hymenolepis prudhoei sp. nov. : hooks. 
FIG. 8. Hymenolepis prudhoei sp. nov. : mature proglottis. 
FIG. 9. Hymenolepis prudhoei sp. nov. : horizontal section of mature proglottis. 



6o 



S. MARKOWSKI 



Hymenolepis prudhoei sp. nov. which in some features may be compared with 
H. Usaccata Fuhrmann, 1906, H. octacantha (Krabbe, 1869) and H. philactes 
Schiller, 1951, differs in the shape and size of the hooks, which in H. bissacata are 
37 (jim long, and in H. octacantha and H. philactes 32-40 (xm and 31-39 (xm 
respectively. Spasski and Spasskaya (1954) give the size of the hooks in H. 
octacantha: 36-38 [xm. 

Although there is some similarity between the new form and the other species 
mentioned, the cirrus-sac is not provided with a saccus accessorius, whilst the 
stylet is smooth and the shape of the testes is also different. 

Some Soviet helminthologists, namely Spasski and Spasskaya (1954), Czaplinski 
(1956), Maksimova (1963), Spasskaya (1966) have erected several new genera by 
breaking down the genus Hymenolepis Weinland (sensu lato). 



10 




II 




FIG. 10. Hymenolepis prudhoei sp. nov. : horizontal section of the cirrus-sac. 
FIG. ii. Hymenolepis prudhoei sp. nov.: male copulatory apparatus. 



PARASITIC WORMS IN THE 'DISCOVERY' COLLECTIONS 



61 



As the erection of these new genera does not seem to have produced a clearly 
understood and concise classification of the species of Hymenolepis (sensu lato), 
Weinland's genus in the sense of Fuhrmann's (1932) is here accepted. 

From the available literature, it seems, that the cestodes recorded from 
Chloephaga picta leucoptera do not include the form described above. Avery (1966) 
gives a list of parasitic worms found in this host, but this list is based entirely upon 
infestation acquired under artificial conditions at Slimbridge, Gloucestershire, 
where the birds are kept in captivity. 



12 






FIG. 12. Hymenolepis prudhoei sp. nov. : transverse section of the segment showing the 

arrangement of longitudinal muscles. 

FIG. 13. Hymenolepis prudhoei sp. nov.: egg. 

FIG. 14. Hymenolepis prudhoei sp. nov. : transverse section of a segment. 



62 



S. MARKOWSKI 



TETRABOTHRIIDAE Fuhrmann, 1908 

9. Tetrabothrius heteroclitus (Diesing, 1850) 

(Figs 15-20) 

Host : Phoebetria fusca : intestine. Locality : Maiviken, West Cumberland Bay, 
South Georgia 14.12.1926. 

Some sixteen complete strobilae and several fragments were collected from the 
intestine of a sooty albatross. 

The length of the strobila is about 18 cm to 20 cm. Its anterior portion is 
narrow and serrated being about 12 cm in length. The posterior part of the strobila 
is 2 mm thick, coiled and shows no distinct segmentation. The mature proglottis 
is 400 [Jim long and i mm broad (Fig. 16). The scolex is 290 (jun long and 330 ^m 
wide, and provided with two 'auriculae' (Fig. 15). The width in that part of the 
scolex is 372 (jon. The suckers are 290 [xm long and 175 [Am across. The longi- 
tudinal muscles form two concentric rings. The inner one is more strongly de- 
veloped (Fig. 19). 



15 




FIG. 15. Tetrabothrius heteroclitus: scolex. 

FIG. 16. Tetrabothrius heteroclitus: mature segment. 

FIG. 17. Tetrabothrius heteroclitus: transverse section of a mature segment. 



PARASITIC WORMS IN THE 'DISCOVERY' COLLECTIONS 63 

The testes, twenty-two to thirty-two, are roundish and situated in the anterior 
and in the posterior part of the segment, as well as in the aporal region of the 
proglottis. They are about 40 [xm in diameter (Fig. 16). The vas deferens forms 
numerous coils. The cirrus-sac is 152 x 150 pm in diameter (Figs 17 and 18). 

The vagina runs ventrally to the cirrus-sac. The ovary is of an irregular shape. 
The egg is 45 X 37 (irn in total diameter. The diameter of the inner membrane 
surrounding the embryo is 33 x 22 [jun. The embryo is 29 x 20 (j.m ; the embryonic 
hooks are 12 [jun long (Fig. 20). From the available literature, it seems that 
Phoebetria fusca is a new host for T. heteroditus. 



18 




20 



FIG. 1 8. Tetrabothrius heteroditus: transverse section of a segment showing cirrus-sac. 
FIG. 19. Tetrabothrius heteroditus : transverse section of a segment showing the arrangement of 

the longitudinal muscles. 
FIG. 20. Tetrabothrius heteroditus: egg. 



64 S. MARKOWSKI 

III. NEMATODA 

HETEROCHEILIDAE (Railliet & Henry, 1915) 
10. Contracaecum osculatum (Rud., 1802) 

Host : Hydrurga leptonyx and Leptonychotes weddelli, intestine. Localities : South 
Orkneys 16.2.1931; Palmer Archipelago 18.1.1935. 

One male and a female specimen were found in the duodenum of H. leptonyx and 
one female in the duodenum of L. weddelli. 

ii. Contracaecum sp. (larvae) 

Host: Chaenocephalus aceratus, liver. Locality: Sandford Bay, South Orkney, 
24.1.1933. 

Mass infestation of the liver of an ice-fish occurred. Because of their larval 
condition the specific determination of the nematodes has not been possible. 

IV. ACANTHOCEPHALA 

POLYMORPHIDAE Meyer, 1931 

12. Corynosoma hamanni (v. Linstow, 1892) 

Host: Notothenia rossi. Locality: South Orkney 17.2.1931. 
Numerous larval stages were found in the mesenteries of the host. Baylis (1929) 
has given a useful description of this species. 



REFERENCES 

AVERY, R. A. 1966. Helminth parasites of wild fowl from Slimbridge, Gloucester. 2. 

Parasites of captive Anatidae. /. Helminth 4 (3) : 269-280. 
BAER, J. G. 1954. Revision taxinomique et etude biologique des Cestodes de la Famille 

de Tetrabothriidae, parasites d'oiseaux de haute mer et de Mammiferes marins. Mem. 

Univ. Neuchdtel 1, pp. 1-121. 
BAYLIS, H. A. 1929. Parasitic Nematoda and Acanthocephala collected in 1925-1927. 

'Discovery' Rep. 1 : 541-560. 
CZAPLINSKI, B. 1956. Hymenolepididae Fuhrmann, 1907 (Cestoda) parasites of some 

domestic and wild Anseriformes in Poland. Acta parasit. pol. 4 : 175-373, figs. 
CZAPLINSKI, B. & RIZHIKOV, K. M. 1964. New data on Parabisaccanthes philactes (Schiller 

1951) Spassky et Reznik, 1963 (Cestoda, Hymenolepididae) from Poland and the Lena 

Delta. Acta parasit. pol. 12 : 363-371. 
DUBININA, M. N. 1953. Cestodes of birds nesting in Western Siberia. Parazit. Sb. 15 : 117- 

233- 

FUHRMANN, O. 1906. Die Hymenolepis-Arten der Vogel. Zentbl. Bakt. ParasitKde (Orig). 

41 : 352-358, 440-452, 39 figs. 
1932. Les Tenias des oiseaux. Mem. Univ. Neuchdtel 8 : 381 pp. 



PARASITIC WORMS IN THE 'DISCOVERY' COLLECTIONS 65 

JOHNSTON, T. H. 1931. New trematodes from the Subantarctic and Antarctic. Aust, J. 
exp. Biol. med. Sci. 8 : 91-98. 

1937. Trematoda. Scient. Rep. Australian, antarct. Exped. 10 : 1-29. 

JOHRI, G. N. 1960. Studies on some Cestodes parasites. IV. On four new species including 

a new genus belonging to the Family Hymenolepididae. Proc. natn. A cad. Sci. India B 

30 : 192-202. 

JOYEUX, CH. & BAER, J. G. 1936. Cestodes. Faune Fr. 30 : 1-613. 
LAP AGE, G. 1961. A list of the parasitic Protozoa, Helminths and Arthropods recorded 

from species of the Family Anatidae (Ducks, Geese and Swans). Parasitology 51 : 1-109. 
MCDONALD, M. E. 1969. Annotated bibliography of Helminths of Waterfowl (Anatidae). 

Spec, scient. Rep. U.S. Fish Wildl. Serv. 125 : 1-333. 
MAKSIMOVA, P. 1963. New species of Cestodes occurring in Kazakhstan swan. Trud. 

Inst. Zool. Alma-Ata 19 : 126-132. 
MARKOWSKI, S. 1952. The Cestodes of seals from the Antarctic. Bull. BY. Mus. not. Hist. 

(Zool.) 1 : 125-150, 21 pis. 

J 955- Cestodes of whales and dolphins from the 'Discovery' collections. 'Discovery' 

Rep. 27 : 377-395- 

NERADOVA, J. 1966. Contribution to the knowledge of the Helminthofauna of domestic 

ducks (Anas platyrhyncha domestica L.) in the western parts of Czechoslovakia. VSst. 

6sl. Spol. Zool. 30 : 247-255. 
PRUDHOE, S. 1969. Cestodes from fish, birds and whales. Rep. B.A.N.Z. antarct. Res. 

Exped. (B) 8 : 171-193- 
RAUSCH, L. R. & FAY, H. F. 1966. Studies on the Helminth fauna of Alaska. XLIV. 

Revision of Ogmogaster jdgerskiold, 1891 with a description of O. pentalineatus sp. n. 

(Trematoda: Notocotylidae) . /. Parasit. 52 : i, 26-38. 
RENNIE, S. & REID, D. 1912. The Cestoda of the Scottish National Antarctic Expedition. 

Rep. Scient. Results Scott, natn. antarct. Exped. Zool. 6 : 243-255, pis. 
SCHILLER, E. L. 1951. Studies on the Helminth fauna of Alaska. VI. The parasites of 

Emperor Goose (Philactes canagica L.) with the description of Hymenolepis philactes n. 

spec. /. Parasit. 37 : 217-229. 

1952. Studies on the Helminth fauna of Alaska. III. Hymenolepis kenaiensis n. 

spec. A Cestode from the Greater Scaup (Ay thy a marila nearctica) with the remarks on 
endemicity. Trans. Am. microsc. Soc. 71 : 146-149. 

SKRYABIN, K. I. & MATEVOSYAN, E. M. 1945. Hymenolepidid Cestodes of domestic and game 

fowl of economic importance. Moscow. 488 pp., Illust. 
SPASSKAYA, L. P. 1966. Cestoda of birds U.S.S.R. Hymenolepididae. Akad. Nauk. Moldav. 

S.S.R. Inst. Zool., 698 pp. 
SPASSKI, A. A. 1963. Hymenolepididae-tapeworms of wild and domestic birds. Part I. 

Principles of Cestodology 2. Akad Nauk S.S.S.R. Moscow, 418 pp. 
SPASSKI, A. A. & SPASSKAYA, L. P. 1954. Construction of the Hymenolepidids system. 

Parasites of birds. Trudy gel'mint. Lab. 7 : 55-119, 27 figs. 
STEFANSKI, W. 1933. Constantin Janicki un distingue Parasitologue Polonais (1876-1932). 

Folia morph. 4 : 220-229, phot. 
SZPOTANSKA, I. 1931. Quelques especes nouvelles ou peu, connues des Hymenolepididae 

Fuhrmann (Cestodes). Pr. zool. pol. panst. Muz. przyr. 9 : 247-266. 



Dr. S. MARKOWSKI 

c/o Department of Zoology 

BRITISH MUSEUM (NATURAL HISTORY) 

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1. KAY, E. ALISON. Marine Molluscs in the Cuming Collection British Museum 
(Natural History) described by William Harper Pease. Pp. 96; 14 Plates. 
1965. (Out of Print.) 3.75. 

2. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and 
Valenciennes. Pp. 180; n Plates, 15 Text-figures. 1967. 4. 

3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure of Mineralogy 
at the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125; 29 Plates, 
77 Text-figures. 1969. 4.50. 

4. HAYNES, J. R. Cardigan Bay recent Foraminifera (Cruises of the R.V. Antur) 
1962-1964. (In press.) 



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MITES\<tF THE GENUS HYPOASPIS 

CANESTRINI, 1884 S.STR. 

AND RELATED FORMS 

(ACARI: MESOSTIGMATA) 

ASSOCIATED WITH BEETLES 



M. COSTA 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 21 No. 4 

LONDON : 1971 



MITES OF THE GENUS HYPOASPIS 
CANESTRINI, 1884 S.STR. AND RELATED FORMS 

(ACARI: MESOSTIGMATA) 
ASSOCIATED WITH BEETLES 




BY 



MICHAEL COSTA 

Kibbutz Mishmar Haemek, Israel 



Pp. 67-98; 101 Text-figures 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 21 No. 4 

LONDON: 1971 



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In 1965 a separate supplementary series of longer 
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This paper is Vol. 21, No. 4 of the Zoological 
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MITES OF THE GENUS HYPOASPIS 
CANESTRINI, 1884 S.STR. AND RELATED FORMS 

(ACARI : MESOSTIGMATA) 
ASSOCIATED WITH BEETLES 

By MICHAEL COSTA 

INTRODUCTION 

RECENTLY Dr C. Athias-Henriot (Laboratoire de Faune du Sol de 1'I.N.R.A., 
Dijon) forwarded to me mite material collected from the scarabaeid beetles Oryctes 
rhinoceros L. and Oryctes monoceros Ol. The beetles and mites were laboratory 
reared and collected by Dr M. J. Stelzer and Mr B. Zelazny (both from U.N./S.P.C. 
Rhinoceros Beetle Project, Apia, Western Samoa). Mr Zelazny informed me that 
the mites feed on the beetles' eggs and may therefore have a possible role in the 
biological control of Oryctes rhinoceros, a main pest of Coco-nut palms. According 
to Mr Zelazny, the mites originally associated with 0. monoceros, imported from the 
Ivory Coast, were even more avid egg feeders on the eggs of 0. rhinoceros than 
the locally collected mites. One batch of mites, ex 0. rhinoceros, was tentatively 
determined as Coleolaelaps rhinocerotis (Ouds.), although I was puzzled by the 
report on their feeding behaviour. It was generally assumed that mites of the genus 
Coleolaelaps Berlese, 1914 are harmless exudate feeders (Grandi, 1925; Vitzthum, 
1940-43). A closer examination of material collected by myself from phytophagous 
scarabaeids in Israel, material collected by Dr M. Remillet (O.R.S.T.O.M., Centre 
d'Adiopodoume, Abidjan, Ivory Coast) and material from melolonthine bettles 
in the U.S.A., showed that 'Coleolaelaps' served actually as a 'dumping ground' 
for a number of different genera of mites associated with beetles. The mites of 
the genus Coleolaelaps Berlese, 1914 have been dealt with in a separate study (Costa 
& Hunter, in press) which has shown that Coleolaelaps is not closely related to 
Hypoaspis Can. The present paper will deal with the genus Hypoaspis s. str. 
and some additional forms. 

The confusion between Coleolaelaps s. str. and Hypoaspis s. str. seems to have 
arisen from the fact that both have long 'wavy' setae on the idiosoma as well as 
macrosetaeon the legs, and both are associated with phytophagous lamellicorn beetles. 
I should like to point out that the 'wavyness' mentioned by many authors is an 
artefact of the preparation, the materials used causing apparently a slight contrac- 
tion of the setal core. In living or alcohol-stored specimens the setae are straight 
or slightly curved. 

Mites of the Hypoaspidinae are generally considered to be the most primitive 
group in the Laelapidae (Vitzthum, 1940-43; Evans, 1958) but the taxonomic 
treatment of Hypoaspis Can. s. lat. remains controversial. This has been shortly 
discussed by Hunter & Costa (in press), who retain at full generic status many of 



7 o M. COSTA 

the subgenera of Hypoaspis which have been recorded, though not used, by Evans 
& Till (1966). The present study supports this view and proposes to show that 
mites of the genus Hypoaspis s. str. are well defined morphologically as well as 
ecologically in their host associations. A close examination of the symbiontic 
mites showed a high degree of host specificity and probably in the past several 
species have been confused with either Hypoaspis krameri Can. (the type species 
of the genus) or Hypoaspis integer Berlese sensu Samsinak, 1960, both of which have 
been also confused with each other. 

Evans & Till (op. cit.) have recently described and figured both sexes of Hypoaspis 
krameri from specimens associated with Lucanus sp. in Great Britain. Their 
description agrees with the details which can be learned from the descriptions of 
H. krameri by G. & R. Canestrini (1881), G. Canestrini (1885) and Berlese (1892), 
making the two undoubtedly congeneric. The host association of the British material 
makes it debatable if this is actually conspecific with the original H. krameri which 
is associated with Oryctes nasicornis L. (compare discussion). 

In view of the present study a redefining of Hypoaspis s. str. seemed to be 
necessary. 

DEPOSITION OF TYPES 

The holotypes are deposited in the British Museum (Nat. Hist.). Paratypes 
will be deposited in The American Museum of Natural History; The Acarina col- 
lection, Department of Entomology, University of Georgia and the author's col- 
lection. 



Hypoaspis Canestrini s. str. 

Hypoaspis Canestrini, 1884, Atti R. 1st. veneto Sci. (6) 2 : 1569; 1885, Acarofauna Ital. part I : 55. 
TYPE: Gamasus krameri Canestrini, 1881. 

FEMALE : Dorsal shield entire, oval, with basically 37 pairs of setae (20 podonotal 
and 17 opisthonotal, fig. i). Setae i2, 54-6 considerably longer than remaining 
setae, setae Z4 extremely long, usually longest idiosomal setae. A tendency exists 
towards diminishing the number of setae, e.g. in Hypoaspis integer Berlese setae 73 
are absent in most specimens; in Hypoaspis phyllognathi sp. n. seta 53 may be absent 
on one or both sides and Hypoaspis remilleti sp. n. lacks setae 33 in all specimens. 
Gnathosoma with six rows of deutosternal denticles, setae Hyp. 3 very long, distinctly 
longer than remaining gnathosomal setae (fig. 7). Sternal shield hexagonal with 
distinct anterior border. Genital shield tongue-shaped with marginally inserted 
genital setae. Paranal setae always longer than postanal seta. Peritreme extends 
anteriorly beyond the margin of coxa I, not attached to dorsal shield. Tarsus II 
with two subterminal stout, blunt, spur-like setae (all and pl\, fig. 4), leg IV with 
macrosetae on femur (adi), genu (ad\ t in some species this seta might be similar 
in length to the remaining setae of the segment) and tarsus (adz, Pdz and pdz, fig. 6). 
Macrosetae are also present on femur II (pdi) and femur III (ad\). Leg chaetotaxy 
as recorded for free-living laelapids (Evans, 1963). 



HYPOASPIS MITES AND BEETLES 71 

MALE: With long slender spermadactyl, curved distally (fig. 14). Holoventral 
shield with 10 pairs of setae in addition to anal setae (various degrees of erosion 
may separate the anal shield completely in Hypoaspis integer). Peritreme anteriorly 
fused with dorsal shield. Leg II with ventral stout, pointed, spine-like setae on 
femur to tarsus (fig. 38). Remaining characters as in female. 

The mites are usually associated with phytophagous scarabaeids, mainly Dyna- 
stinae. 



Hypoaspis neokrameri sp. n. 

FEMALE : Dorsum covered by single dorsal shield (735 (xm long and 445 |xm wide) 
with 37 pairs of setae. Podonotal setae iz, si, 54-6 are elongate and distinctly 
longer than the remaining podonotal setae (fig. i). The longest dorsal setae are 
Z4 (220 (xm), setae J5 straight and short (30 (xm). The shield is nearly devoid 
of ornamentation. Tectum (fig. 3) triangular with denticulate proximal margins. 

Tritosternum normal with pilose laciniae. Sternal shield (170 |xm long and 
150 (Am wide at St2) well ornamented, posterior margin slightly convex and irregular 
(fig. 2). Sternal setae long, reaching to or beyond the bases of consecutive setae. 
Genital shield (distance between genital setae 105 [xm) ornamented and tongue- 
shaped, metapodal shields narrow. Paranal setae distinctly longer than postanal 
seta. Peritreme extending anteriorly slightly beyond the middle of coxa I, free 
anteriorly and posteriorly. 

Gnathosoma (fig. 7) with well-sclerotized corniculi and fimbriate internal malae. 
Six rows of deutosternal denticles (8-14 per row). Movable digit of chelicera 
(fig. 5) bidentate, fixed digit with one stout tooth and about ten small denticles. 

The approximate lengths of the legs (excluding pretarsi) are: I 660 (xm; II 
540 (Jim ; III 600 (xm ; IV 850 (xm. Tarsus II (fig. 4) with two blunt distal spines, 
leg IV with macrosetae on the femur (220 (xm), genu and tarsus (fig. 6). Macrosetae 
present also on femur II (150 (xm) and III (no (xm). Leg chaetotaxy normal. 

MALE: Unknown. 

DIFFERENTIAL DIAGNOSIS : H. neokrameri sp. n. can be separated from other 
species of the complex by its long sternal shield which is longer than wide. Associ- 
ated with Oryctes nasicornis L. (Scarabaeidae : Dynastinae). 

MATERIAL: Holotype: , ex Oryctes nasicornis L., Tivon, Israel, May 25, 1965, 
coll. M. Costa. Paratypes: 5$?, ibid. 



Hypoaspis pentodoni sp. n. 

FEMALE: Dorsum covered by a single dorsal shield (680 (xm long and 390 |xm 
wide) with 37 pairs of setae. Podonotal setae 12, 54-6 are elongate and distinctly 
longer than the remaining podonotal setae (fig. 8). The longest dorsal setae are 
Z4 (220 (xm). The shield is nearly devoid of ornamentation, sites of muscle attach- 
ments are striated. Tectum (fig. 12) triangular with denticulate margins. 



M. COSTA 




FIGS 1-7. Hypoaspis neokrameri sp. n., female. Fig. i. Dorsum. Fig. 2. Venter. 
Fig. 3. Tectum. Fig. 4. Tarsus II. Fig. 5. Chelicera. Fig. 6. Leg IV. Fig. 7. 
Gnathosoma, ventral view. 



HYPOASPIS MITES AND BEETLES 



73 



Tritosternum normal with pilose laciniae. Sternal shield wider than long (125 [xm 
long and 150 ^m wide at St2), its posterior margin is nearly straight with two small 
but characteristic projections (fig. 9). Sternal setae long, extending beyond the 




13 



FIGS 8-15. Hypoaspis pentodoni sp. n., female. Fig. 8. Dorsum. Fig. 9. Venter. 
Fig. 10. Leg IV. Fig. n. Tarsus II. Fig. 12. Tectum. Fig. 13. Chelicera. 
Fig. 15. Gnathosoma, ventral view. Male. Fig. 14. Chelicera. 



74 M. COSTA 

bases of consecutive setae. The ornamented genital shield is tongue shaped (dis- 
tance between genital setae no (xm). Metapodal shields narrow. Postanal seta 
distinctly shorter than paranal setae. The peritreme extends anteriorly beyond the 
middle of coxa I. 

Gnathosoma (fig. 15) with well-sclerotized corniculi and fimbriate internal malae, 
six rows of tiny deutosternal denticles. Movable digit of chelicera (fig. 13) bidentate, 
fixed digit with small denticles. 

The approximate lengths of the legs (excluding pretarsi) are: I 650 (xm; II 
540 (xm; III 530 fxm; IV 750 [xm. Tarsus II (fig. n) with two blunt distal 
spines. Leg IV (fig. 10) with macrosetae on femur (210 (Am), genu and tarsus 
(^3165 (Jim). 

MALE : The single male specimen is smaller than the female (dorsal shield 580 (j.m 
long and 330 [xm wide). Venter covered by well-ornamented holo ventral shield 
with 10 pairs of setae in addition to the anal setae. Chelicera (fig. 14) with slender, 
distally curved, spermadactyl. Remaining characteristics as in female, except 
peritreme which is attached anteriorly to the dorsal shield. 

DIFFERENTIAL DIAGNOSIS: This species can be separated from H. neokrameri sp. n. 
by its short sternal shield and from H. phyllognathi sp. n. by its shorter dorsal 
setae. Associated with Pentodon bispinosus Kiist (Scarabaeidae : Dynastinae). 

MATERIAL: Holotype: 9- ex Pentodon bispinosus Kiist, Mishmar Haemek, Israel, 
Sept. 21, 1962 coll. M. Costa. Paratypes: all from the same host and locality: 
i<$, i$, May 19, 1965; 5$$, Sept. 21, 1962; 6.?, Sept. 12, 1966; i$, Nov. 8, 1965. 

Hypoaspis phyllognathi sp. n. 

FEMALE : Dorsum covered by single dorsal shield (840 {xm long and 470 [xm wide) 
which is ornamented with small polygons mainly in the posterior area. Regularly 
37 pairs of simple setae are inserted on the shield, but in several specimens setae 
53 are absent on one or both sides. Podonotal setae i2 and 54-6 are distinctly 
longer than the remaining podonotal setae. The longest dorsal setae are Z4 
(320 (xm), setae J5 straight and short (30 fxm). The distribution and relative 
lengths of the setae are shown in fig. 16. Tectum (fig. 21) triangular, proximal 
margins deeply denticulate. 

Tritosternum normal with pilose laciniae. Sternal shield (145 (xm long and 
155 [xm wide at St2) well sclerotized, only faintly ornamented. Sternal setae 
very long, St2 extends beyond the posterior margin of the shield (fig. 17). The 
tongue-shaped genital shield (distance between genital setae (125 (xm) has nearly 
parallel sides and is well ornamented. The anal shield is rounded anteriorly, the 
postanal seta is distinctly shorter than the paranal setae. The anal shield is flanked 
posteriorly by a pair of long setae. Metapodal shields small, irregular in shape. 
The peritreme extends anteriorly to the anterior margin of coxa I, it is free both an- 
teriorly and posteriorly. 

Gnathosoma (fig. 18) with well-sclerotized corniculi and fimbriate internal malae, 
with six rows of minute deutosternal denticles. Movable digit of chelicera (fig. 22) 



HYPOASPIS MITES AND BEETLES 



75 




FIGS 16-22. Hypoaspis phyllognathisp.n., female. Fig. 16. Dorsum. Fig. 17. Venter. 
Fig. 18. Gnathosoma, ventral view. Fig. 19. Tarsus II. Fig. 20. Leg IV. Fig. 21. 
Tectum. Fig. 22. Chelicera. 



76 M. COSTA 

bidentate, fixed digit with 8 small denticles in addition to a large tooth which is 
associated with the pilus dentilis. 

The approximate lengths of the legs (excluding pretarsi) are: I 730 [xm; II 
570 (xm; III 600 (Am; IV 870 jjun. Tarsus II (fig. 19) with two dorsal distal 
blunt spines, leg IV (fig. 20) with macrosetae on the femur (300 |xm), genu (210 [xm) 
and tarsus. Macrosetae are also present on femora II and III, leg chaetotaxy 
normal. 

MALE: Unknown. 

DIFFERENTIAL DIAGNOSIS: This species can be recognized by its long dorsal 
setae, its large size and by the long seta adi (210 [xm) on genu IV. Associated 
with Phyllognathus silenus F. (Scarabaeidae : Dynastinae) . 

MATERIAL: Holotype: $, ex Phyllognathus silenus F., Bardawil, Northern Sinai, 
April 10, 1968, beetle coll. H. Sandier. Paratypes: 13 <j><j>, same data; 6 $<j>, Carmia, 
Israel, Nov. 16, 1966; I -, Tivon, Israel, May 6, 1966; 2 .., Ein Yahav, Israel, 
May i, 1968. 

Hypoaspis integer Berlese, 1911 sensu Samsinak, 1960 

Laelaps (Hypoaspis} integer Berlese, 1911, Redia 7 : 186. 

? 'Coleolaelaps integer Willmann, 1935, Bull. Mus. R. Hist. nat. Belg. n : 23-25, figs 14-6. 
Coleolaelaps integer (male) Samsinak, 1960, Cas. dsl. Spol. ent. 57 (3) : 280-82, figs 1-6. 

FEMALE : Dorsum covered by single dorsal shield (820 pan long and 430 [xm wide) 
with 36 37 pairs of setae. Of 29 investigated specimens, setae Z3 were completely 
absent in 19 specimens, unilaterally present in 8 and bilaterally present in 2 speci- 
mens only. Podonotal setae 12, si, 54-6 are elongate, the longest dorsal setae are 
Z4 (280 |xm) and a pair of postero-marginal setae (200 |xm) which are inserted on 
the soft integument. The shield is devoid of distinct ornamentation, the distri- 
bution and relative lengths -of the setae are shown in fig. 23. Tectum (fig. 25) 
triangular with denticulate margin. 

Tritosternum normal with pilose laciniae. Sternal shield (150 [xm long and 
150 [xm wide at St2) ornamented with short sternal setae which do not reach the 
bases of the consecutive setae (fig. 24). Metasternal setae inserted on integument. 
The large genital shield (distance between genital setae 130 jxm) expands slightly 
beyond the genital setae and is ornamented. Metapodal shields narrow, kidney- 
shaped. Anal shield with semicircular anterior border, postanal seta distinctly 
shorter than paranal setae. The anal shield is flanked posteriorly by a pair of long 
(300 (xm) setae. The narrow peritreme extends anteriorly to the middle of coxa I. 

Gnathosoma (fig. 29) with well-sclerotized corniculi and fimbriate internal malae, 
six rows of tiny deutosternal denticles (about 12 per row). Movable digit of chelicera 
(fig. 27) bidentate, fixed digit with one large tooth and about 10 sub-equal small 
denticles. 

The approximate lengths of the legs (excluding pretarsi) are: I 680 |xm; II 
620 (xm; III 580 (xm; IV 960 [xm. Tarsus II (fig. 28) with two dorsal distal 
blunt spines, with ventral setae markedly stouter than the dorsal setae. Leg IV 



HYPOASPIS MITES AND BEETLES 



77 



(fig. 26) with macrosetae on femur (360 ^m), on the genu (ad-\_ 250 (xm, pdi 
180 (jon) and tarsus; macrosetae are also present on femur II (270 fxm) and III 
(260 (Am) . Leg chaetotaxy normal for the genus. 




27 



FIGS 23-29. Hypoaspis integer Berlese, female. Fig. 23. Dorsum. Fig. 24. Venter. 
Fig. 25. Tectum. Fig. 26. Leg IV. Fig. 27. Chelicera. Fig. 28. Tarsus II. 
Fig. 29. Gnathosoma, ventral view. 



78 M. COSTA 

MALE: The male has been described by Samsinak (1960). The ventral sclerotiza- 
tion is extremely variable, out of 14 male specimens only one had a complete holo- 
ventral shield, in one specimen this was eroded but the anal shield was still broadly 
connected to the ventral shield and in 12 specimens, with variously shaped genito- 
ventral shields, the anal shield was completely separate. In the males the cor- 
responding macrosetae are longer than in the females. 

DIFFERENTIAL DIAGNOSIS: The long macrosetae on femora II and III, as well as 
the two macrosetae on genu IV are good diagnostic characters for this species. 
Associated with Oryctes nasicornis L. (Scarabaeidae : Dynastinae) and Polyphylla 
fullo L. (Scarab.: Melolonthinae). 

NOTES: The original association seems to be with 0. nasicornis, the association 
with P. fullo appears to be secondary (vide Costa & Hunter, in press). The speci- 
mens ex P. fullo (the beetles were kindly loaned by the American Museum of Natural 
History) were collected from beneath the elytra by methods described by Costa 
& Hunter (op. cit.). 

MATERIAL: 2 $$, 28 $$, ex Oryctes nasicornis, Bohemia, Liblice, July 16, 1960, 
coll. and det. K. Samsinak; 5 <$J, 53 ?$ Polyphylla fullo, S. Russia; 3 <$<$, 41 $?, 
ibid., Prussia; 4c&? 56 $$, ibid., Germany, 1897. 



Hypoaspis rhinocerotis Oudemans, 1925 

Hypoaspis rhinocerotis Oudemans, 1925, Ent. Ber., Amsl. 7 (146) : 30. 

Coleolaelaps rhinocerotis Oudemans, 1927, Zoo/. Meded. Leiden 10 (4) : 189-193, figs 8-15. 

FEMALE: Dorsum covered by a single dorsal shield (770-830 (im long and 470- 
540 (Jim wide), with 37 pairs of simple setae. Setae si subequal in length with vertical 
setae (ii) ; the longest podonotal setae being i2, 54-6 (35 being shorter than either 
54 or s6). The central dorsal setae, especially on the opisthonotum are short and 
do not extend to the bases of the consecutive setae. Setae Z4 are the longest 
dorsal setae (300 (Am), setae J5 are short (30 [xm). The distribution and the relative 
lengths of the setae are shown in fig. 30. Tectum (fig. 32) triangular with denticulate 
margin. 

Tritosternum normal with well developed laciniae. Sternal shield (170 fun 
long and 185 (Jim wide at St2) only slightly ornamented. Sti distinctly shorter 
than St2~4 which are long and extend markedly beyond the bases of the consecutive 
setae (fig. 31). The posterior margin of the shield is irregular. Genital shield 
(distance between genital setae 130 (i,m) tongue-shaped and nearly devoid of orna- 
mentation. Metapodal shields narrow and irregular. Anal shield small, postanal 
seta distinctly shorter than paranal setae. Peritreme extends anteriorly slightly 
beyond the middle of coxa I, it is free anteriorly and posteriorly. 

Gnathosoma (fig. 36) with well-sclerotized corniculi and fimbriate internal malae, 
six rows of deutosternal denticles (12-18 per row). Chelicera (fig. 35) with bidentate 
movable digit with a large distance between the two teeth, fixed digit slightly curved 
and sickle shaped with one stout tooth and about 12 small denticles. 



HYPOASPIS MITES AND BEETLES 



79 



The approximate lengths of the legs (excluding pretarsi) are: I 750 [xm; II 
660 (Am; III 700 [xm; IV 950 (j.m. Tarsus II (fig. 34) with two dorsal blunt 
spines, its ventral setae being much stouter than its dorsal setae. Leg IV (fig. 33) 
with macrosetae on femur (270 fxm) and tarsus only, ad\ of the genu being only 
slightly longer than the remaining setae on the segment. Macrosetae are present 
also on femora II and III. Leg chaetotaxy normal. 




FIGS 30-36. Hypoaspis rhinocerotis Oudemans, female. Fig. 30. Dorsal shield. Fig. 31. 
Venter. Fig. 32. Tectum. Fig. 33. Leg IV. Fig. 34. Tarsus II. Fig. 35. Cheli- 
cera. Fig. 36. Gnathosoma, ventral view. 



8o 



M. COSTA 



MALE: Dorsal shield smaller (740 [j,m long and 480 (im wide) than in female, 
with the same chaetotaxy. The venter (fig. 37) is covered by a well- ornamented 
holoventral shield with 10 pairs of setae in addition to the regular anal setae. 
Remaining ventral features as in female. 



500, 




FIGS 37-43. Hypoaspis rhinocerotis Oudemans. Male, Fig. 37. Venter. Fig 38. Leg 
II. Fig. 40. Chelicera. Deutonymph. Fig. 39. Dorsal shield. Fig. 43. Venter. 
Protonymph. Fig. 41. Dorsum. Fig. 42. Venter. 



HYPOASPIS MITES AND BEETLES 81 

Movable digit of chelicera (fig. 40) monodentate with long slender spermadactyl 
which is distally curved. Fixed digit with about 4 teeth. Leg chaetotaxy as in 
female, leg II (fig. 38) with several stout pointed spine-like setae on femur to tarsus, 
similar to the condition found in H. krameri (as figured by Evans & Till, 1966). 

DEUTONYMPH : Dorsal shield (670 [xm long and 380 [xm wide) deeply incised later- 
ally, chaetotaxy and other features as in female although several central setae 
are longer (fig. 39). Sternal shield (fig. 43) extends only slightly beyond the posterior 
margin of coxa IV. A number of small platelets are present on the integument. 
Remaining characters as in female. 

PROTONYMPH : The idiosoma (660 (xm long) is covered by two dorsal shields and 
three pairs of platelets in the mesonotal region. The podonotal shield (365 |xm 
long and 325 [xm wide) with n pairs of long setae all of which extend beyond the 
bases of consecutive setae. The opisthonotal shield with 8 pairs of setae, 85, 
Z4~5 and J5 having the same relative lengths as in the adult. The distribution and 
relative lengths of the setae are shown in fig. 41. The venter (fig. 42) with a sternal 
shield with 3 pairs of setae. Peritremes rudimentary. 

DIFFERENTIAL DIAGNOSIS: The short central dorsal setae and the large sickle- 
shaped chelicera separate H. rhinocerotis from the preceding species. Associated 
with Oryctes rhinoceros L. (Scarabaeidae : Dynastinae). 

MATERIAL: Numerous specimens, ex Oryctes rhinoceros, Apia, W. Samoa, July, 
1969; additional specimens from eggs of 0. rhinoceros and laboratory cultures. 
All the specimens were made available through the courtesy of Mr B. Zelazny and 
Dr M. J. Stelzer. 

Hypoaspis athiasae sp. n. 

FEMALE: Dorsal shield (770-820 [xm long and 490-510 fxm wide) covers most of 
the dorsum, with 37 pairs of setae. The shield is slightly ornamented, mainly on 
its posterior part. Setae i2, 54-6 are the longest podonotal setae, 55 being markedly 
shorter than either 54 or s6. Setae Z4 are the longest dorsal setae (280 |xm), J5 
are short (30 [xm). The distribution and the relative lengths of the setae are shown 
in fig. 44. Tectum (fig. 46) triangular with denticulate margin. 

Tritosternum with well-developed laciniae. Sternal shield (195 [xm long and 
175 [Jim wide at St2) without apparent ornamentation, with long sternal setae 
which extend beyond the base of the consecutive setae. Posterior margin of shield 
nearly straight and slightly irregular. Genital shield (distance between genital 
setae 120 (xm) tongue-shaped and well ornamented. Narrow, kidney-shaped meta- 
podal shields. The wide peritreme extends anteriorly to the anterior margins of 
coxa I. The postanal seta is shorter than the paranal setae. 

Gnathosoma (fig. 50) with well-sclerotized corniculi and fimbriate internal malae, 
with six rows of deutosternal denticles (10-14 per row). Movable digit of chelicera 
(fig. 48) bidentate, fixed digit with about 14 small denticles in addition to a larger 
tooth which is associated with the pilus dentilis. 



82 



M. COSTA 



The approximate lengths of the legs (excluding pretarsi) are: I 840 |xm; II 
675 fxm; III 710 (xm; IV 800 {xm. Tarsus II (fig. 47) with two dorsal distal 
blunt spines. Leg IV (fig. 49) with macrosetae on femur (280 fxm) and tarsus 




FIGS 44-50. Hypoaspis athiasae sp. n., female. Fig. 44. Dorsal shield. Fig. 45. 
Venter. Fig. 46. Tectum. Fig. 47. Tarsus II. Fig. 48. Chelicera. Fig. 49. Leg 
IV. Fig. 50. Gnathosoma, ventral view. 



HYPOASPIS MITES AND BEETLES 



only, adi of the germ being only slightly longer than the remaining setae of the 
segment. Macrosetae also present on femora II and III. 

MALE: The dorsal shield (815 [Ain long and 485 ^m wide) and chaetotaxy as in 
female. Ventrally the idiosoma is covered by a well-ornamented holo ventral 




FIGS 51-57. Hypoaspis athiasae sp. n. Male, Fig. 51. Venter. Fig. 52. Leg II. Fig. 
54. Chelicera. Deutonymph, Fig. 53. Dorsal shield. Fig. 57. Venter. Proto- 
nymph, Fig. 55. Dorsum. Fig. 56. Venter. 



84 M. COSTA 

shield, usually with 10 pairs of setae in addition to the regular anal setae (fig. 51). 
The extent and outlines of the shield may vary asymetrically and with it the chaeto- 
taxy. The chelicera (fig. 54) with monodentate movable digit which bears a long 
slender spermadactyl, slightly curved distally. Leg II with pointed spine-like 
setae on femur to tarsus (fig. 52), similar to the condition in H. krameri (Evans & 
Till, op. cit.). 

DEUTONYMPH : Dorsal shield (715 [xm long and 400 (xm wide) deeply incised later- 
ally. Chaetotaxy similar to that of the female, central setae (mainly i series) notice- 
ably long. The shield is faintly ornamented mainly on its posterior portion. The 
distribution and relative lengths of the setae is shown in fig. 53. The anal shield 
is rounded and the postanal seta is markedly shorter than the paranal setae (fig. 57). 

PROTONYMPH: The idiosoma (530 [xm long) is covered by two dorsal shields and 
three pairs of platelets in the mesonotal region. The podonotal shield (350 {xm 
long and 290 (xm wide) with n pairs of long setae. The opisthonotal shield (120 pun 
long and 170 (xm wide) with 8 pairs of setae. Setae 85, Z4~5 and J5 have the same 
relative lengths as in the adult. The distribution and the relative lengths of the 
setae are shown in fig. 55. The venter (fig. 56) with small sternal shield which has 
a very indistinct anterior margin. 

DIFFERENTIAL DIAGNOSIS: This species is closely related to H. rhinocerotis from 
which it can be separated mainly by its longer central dorsal setae, a character 
which is even more conspicuous in the deutonymph. 

NOTES : This species has been collected in large numbers (by Dr C. Athias-Henriot) 
from soil as well as from eggs of 0. monoceros in the Ivory Coast. It is well known 
that many dynastine beetles emerge from their puparia during a few days of the 
year only, usually for a short time before dusk. It may well be that the possibility 
to collect a large number of the mites from the soil happened after a mass emergence 
of the beetles which are probably their normal host. 

MATERIAL: Holotype: $, ex humid soil (Aerodrome), Ivory Coast, July 18, 1969, 
coll. C. Athias-Henriot. Paratypes: i $ and numerous female specimens ibid. ; 
5 $$ Oryctes monoceros, Ivory Coast, 1969. 



Hypoaspis dubius sp. n. 

FEMALE: Dorsum covered by a single dorsal shield (640 [xm long and 380 {xm 
wide), with 37 pairs of simple setae. The shield is finely ornamented mainly on its 
posterior part. Setae 12, si and 54-6 are longer than the remaining podonotal 
setae, the longest dorsal setae are Z4 (130 (xm), the shortest J5 (25 |xm). Nearly 
all the central setae are long and extend beyond the base (or the horizontal level 
of the bases) of the consecutive setae. The distribution and the relative lengths 
of the setae are shown in fig. 58. Tectum (fig. 61) triangular with denticulate 
margin. 



HYPOASPIS MITES AND BEETLES 



Tritosternum normal with pilose laciniae. Sternal shield (135 |xm long and 140 |j.m 
wide at St2) faintly ornamented mainly in its anterior and lateral parts, posterior 
margin slightly concave to nearly straight. Sti shorter than remaining sternal 
setae which are long and extend beyond the base of consecutive setae. The tongue- 




FIGS 58-64. Hypoaspis dubius sp. n., female. Fig. 58. Dorsal shield. Fig. 59. Venter. 
Fig. 60. Leg IV. Fig. 61. Tectum. Fig. 62. Tarsus II. Fig. 63. Chelicera. 
Fig. 64. Gnathosoma, ventral view. 



86 M. COSTA 

shaped genital shield (distance between genital setae 105 (xm) is faintly ornamented 
(fig. 59). Metapodal shields narrow and elongate. The postanal seta is shorter 
than the paranal setae. The peritreme extends anteriorly slightly beyond the middle 
of coxa I. 

Gnathosoma (fig. 64) with well-sclerotized corniculi and fimbriate inner malae, 
six rows of deutosternal denticles (14-18 per row). Movable digit of chelicera 
(fig. 63) bidentate, fixed digit with about 8 small denticles in addition to one larger 
tooth. 

The approximate lengths of the legs (excluding pretarsi) are: I 640 (xm; II 
490 (Jim; III 490 {xm; IV 690 (xm. Tarsus II (fig. 62) with two dorsal distal 
blunt spines. Leg IV (fig. 60) with macrosetae on the femur (145 |xm) and tarsus 
only. Macrosetae are present also on femora II and III. 

MALE: Unknown. 

DIFFERENTIAL DIAGNOSIS: This species is characterized by its small size and 
short Z4 and ad\ of femur IV. 

NOTES : The status of this species is uncertain and it might be a hybrid. Originally 
it was introduced from the Ivory Coast, ex Oryctes monoceros. It has become 
established in W. Samoa and occurred together with H. rhinocerotis on a field- 
collected 0. rhinoceros (coll. Dr M. J. Stelzer). In view of the fact that Hypoaspis 
athiasae sp. n. has also been collected from 0. monoceros, a misdetermination of the 
African host cannot be ruled out, since over a dozen species of Oryctes exist in central 
and western Africa. 

MATERIAL : Holotype : $, laboratory reared, Apia, W. Samoa, July 1969, laboratory 
colony started from specimens ex Oryctes monoceros, Ivory Coast. Paratypes: 
many ibid. ; 6 -$, 0. rhinoceros, Apia, W. Samoa, Sept. 1969, coll. Dr M. J. Stelzer. 

Hypoaspis remilleti sp. n. 

FEMALE: Dorsum covered by single dorsal shield (870 |xm long and 560 fxm 
wide) leaving a wide strip of uncovered integument. The shield bears 36 pairs of 
setae (53 missing, compare Hypoaspis phyllognathi sp. n.) of three different types: 
minute setae (about 20 [xrn long: 13-5, Z2-3, J series px2~3, Zi-3 and Si) ; extremely 
long setae (i2, 54-6 and Z4 which are 400 ^m long) and 'normal' marginal setae. 
The posterior elongate integumental setae are only 180 jxm long. The distribution 
and the relative lengths of the setae are shown in fig. 65. Tectum (fig. 66) triangular 
with denticulate margin. 

Tritosternum normal with pilose laciniae. Sternal shield (160 (xm long and 170 [xm 
wide at St2) ornamented, with irregular posterior margin (fig. 67). The sternal 
setae are long, extending beyond the bases of the consecutive setae. The well- 
ornamented genital shield (distance between genital setae 150 (xm) expands slightly 
beyond coxae IV. Anal shield with round anterior margin, postanal seta shorter 
than paranal setae. The peritreme extends anteriorly to or slightly beyond the 
anterior margin of coxa I, it is free both anteriorly and posteriorly. 



HYPOASPIS MITES AND BEETLES 



Gnathosoma (fig. 71) with well-sclerotized corniculi and fimbriate internal malae, 
six rows of minute deutosternal denticles. Movable digit of chelicera (fig. 68) 
bidentate, fixed digit with about 12 small denticles in addition to one larger tooth. 




FIGS 65-71. Hypoaspis remilleti sp. n., female. Fig. 65. Dorsum. Fig. 66. Tectum. 
Fig. 67. Venter. Fig. 68. Chelicera. Fig. 69. Tarsus II. Fig. 70. Leg IV. 
Fig. 71. Gnathosoma, ventral view. 



88 M. COSTA 

The approximate lengths of the legs (excluding pretarsi) are: I 890 pan; II 
740 pan; III 800 pan; IV noo pan. Tarsus II (fig. 69) with two blunt dorsal 
subterminal spines. Leg IV (fig. 70) with macrosetae on the femur (380 pan) 
and tarsus only. Leg chaetotaxy normal. 

MALE : Dorsal shield as in female. The venter is covered by a holoventral shield 
with 10 pairs of setae in addition to the regular anal setae. Leg II with pointed 
spine-like ventral setae on femur to tarsus. Chelicera with slender spermodactyl, 
distally curved. 

DIFFERENTIAL DIAGNOSIS : This species can be easily recognized by its very short 
central setae and its long antero-lateral setae. It differs from closely related 
'Coleolaelaps' proximus Cooreman, 1948 mainly in the shape of the sternal shield 
which is markedly longer than wide in C. proximus. Associated with Heteroligus 
meles Billb. (Scarabaeidae : Dynastinae). 

MATERIAL: Holotype: ., ex Heteroligus meles Billb., nr. Abidjan, Ivory Coast, 
1969, coll. Dr M. Remillet. Paratypes: numerous specimens including males with 
the same data. 

Lucanaspis gen. n. 

General facies of female as in Hypoaspis with the following differences: Thirty- 
three pairs of dorsal setae of which i2, 54 and Z4 are very long. Sternal shield 
markedly wider than long. Legs stumpy and shorter than length of dorsal shield. 
Tarsus II with two subterminal stout and pointed setae. 

MALE: Unknown. 

Lucanaspis brachypedes sp. n. 

FEMALE: Dorsum covered by a single dorsal shield (545 pan long and 360 pan 
wide), devoid of ornamentation, with 33 pairs of simple setae. The podonotal 
setae (17 pairs) are very short (10-15 pan) with the exception of ii and si (85 pan), 
12, 54 (120 pan), 55 and s6 (60 pan). The opisthonotal setae (16 pairs) are short 
with the exception of Z4 (155 pan) which are the longest dorsal setae. The distri- 
bution and the relative lengths of the setae are shown in fig. 72. Tectum (fig. 78) 
triangular with denticulate margins. 

Tritosternum normal with pilose laciniae. Sternal shield (75 pan long and 130 pan 
wide at St2) with distinct anterior concave margin and slightly irregular concave 
posterior margin. The shield is devoid of distinct ornamentation. The first sternal 
pores are very large and close to each other. Metasternal setae inserted on the 
integument, the associated pore may be incorporated into the sternal shield (fig. 73). 
Genital shield (distance between genital setae 95 pan) broad and tongue-shaped. 
Anterior margin of anal shield semicircular, paranal setae slightly longer than postanal 
seta. The shield is flanked by two pairs of slightly longer and stouter ventral 
setae. Metapodal shields elongate, oval. The peritreme extends anteriorly 
nearly to the anterior margin of coxa I, it is free both anteriorly and posteriorly. 



HYPOASPIS MITES AND BEETLES 



Gnathosoma (fig. 75) with well-sclerotized corniculi and fimbriate internal malae, 
with six rows of tiny deutosternal denticles. Movable digit of the chelicera (fig. 74) 
bidentate, fixed digit with three medium sized distal teeth and about seven small 
proximal denticles. 




74 



76 



FIGS 72-78. Lucanaspis brachypedes gen. n., sp. n., female. Fig. 72. Dorsal shield. 
Fig- 73- Venter. Fig. 74. Chelicera. Fig. 75. Gnathosoma, ventral view. Fig. 
76. Tarsus II. Fig. 77. Leg IV. Fig. 78. Tectum. 



go M. COSTA 

The approximate lengths of the legs (excluding pretarsi) are: I 520 (xm; II 
450 (jim; III 440 (Jim; IV 540 (Jim. Leg chaetotaxy normal as in free-living 
laelapids. Macrosetae are present on femora II-IV (on femur IV 165 [xm) and 
tarsus IV (fig. 77). Tarsus II (fig. 76) with two subterminal pointed stout setae. 

MATERIAL: Holotype: <j>, ex lucanid beetle, nr. Abidjan, Ivory Coast, Oct. 1969, 
coll. Dr M. Remillet. Paratypes: 8 $?, same data. 

Dynastaspis gen. n. 

General facies of female as in Hypoaspis with the following differences: 32 pairs 
of dorsal setae (18 podonotal and 14 opisthonotal) with 54-6 and Z^ very long and 
'wavy'. Tectum with nondenticulate margin. Four macrosetae of a different 
homology on tarsus IV: adz, alz, Ph and ^^3 (instead oipdz, pd$ and adz in Hypoaspis 
s. str.), pdz being proximally inserted to adz (as distinct from being at the same level 
in Hypoaspis s. str.). Tarsus II with two subterminal pointed stout setae. Orna- 
mentation of genital shield different from that found in Hypoaspis. 

MALE: Unknown. 

Dynastaspis walhallae sp. n. 

FEMALE : A single dorsal shield (1030 (xm long and 580 jxm wide) covers the dorsum 
incompletely, faintly ornamented with a scale-like pattern (fig. 86), with 32 pairs 
of simple setae (fig. 79). Eighteen pairs of podonotal setae, Z3 absent, 57 on the 
integument and 14 pairs of opisthonotal setae (84 and px2~3 absent). Medial 
setae mainly short (e.g. 14 45 (Jim), 54-6 long and 'wavy', Z^ being the longest 
dorsal setae (400 (xm). Tectum (fig. 83) with nondenticulate margin and broad 
median projection. 

Tritosternum normal with 'pilose laciniae. Sternal shield (140 (jim long and 
195 (xm wide at St2), well ornamented with concave anterior margin and deeply 
concave posterior margin (fig. 80, 8ia-c). Sternal setae long, St2 extending beyond 
posterior margin of shield. The shape of the sternal shield is variable, mainly 
in the lateral outlines (figs 8ia-c, in 8ic one Sti completely missing). Genital 
shield (distance between genital setae 155 [xm) tongue-shaped, with very distinct 
ornamentation which is completely different from that found in species of Hypoaspis 
s. str. Metapodal shields small and irregular, several small platelets present at 
the same level. Anal shield nearly triangular in shape, postanal seta shorter than 
the paranal setae. The peritreme extends anteriorly slightly beyond the posterior 
margin of coxa I, it is free anteriorly and posteriorly. 

Gnathosoma (fig. 87) with well-sclerotized corniculi and fimbriate internal malae, 
with six rows of deutosternal teeth (10-14 P er row). Movable digit of the chelicera 
(fig. 85) bidentate, fixed digit with about eight denticles of different sizes and shapes. 

The approximate lengths of the legs (excluding pretarsi) are: I 1010 jxm; II 
850 (xm; III 870 [xm; IV 1190 jxm. Tarsus II (fig. 84) with pointed setae only, 
leg IV (fig. 82) with macrosetae on femur (400 (xm) and four macrosetae (adz, <*h, 



HYPOASPIS MITES AND BEETLES 91 

plz, pdz) on the tarsus with pdz proximally inserted. Otherwise leg chaetotaxy 
normal. 




FIGS 79-87. Dynastaspis walhallae gen. n., sp. n., female. Fig. 79. Dorsum. Fig. 80. 
Venter. Fig. 8ia-c. Variations in shape of sternal shield. Fig. 82. Leg IV. Fig. 
83. Tectum. Fig. 84. Tarsus II. Fig. 85. Chelicera. Fig. 86. Ornamentation of 
dorsal shield. Fig. 87. Gnathosoma, ventral view. 



92 M. COSTA 

MATERIAL: Holotype: -, ex larvae of Dynastes tytius Linn. (Scarabaeidae : 
Dynastinae), dead Black Locust Tree, Walhalla, South Carolina, August 1969, 
coll. M. Palmer. Paratypes: 8 .-, same data as type. 

NOTES: The occurrence of this species on larvae of Dynastes tytius is rather sur- 
prising as Sikora (1968) reported a different species of 'Coleolaelaps' from the adult 
of D. tytius. It may be that the host association of D. walhallae is accidental, the 
true host being another woodboring beetle. 



Angosomaspis gen. n. 

FEMALE: Single dorsal shield with a large number (over 100) of minute (15- 
20 (jun) setae, ventral integument with about 30 pairs of long, very attenuated 
whip-like setae. Macroseta of femur IV exceedingly long, extending beyond middle 
of tarsus IV, only one macroseta (pd%) on tarsus IV. Long macroseta also on femora 
II and III. Tarsus II with two subterminal spine-like setae which are originally 
pointed, though usually the tip is broken off, leaving a sharp (not blunt) end. 
Fixed digit of chelicera with about a dozen sharp, needle-shaped denticles proximal 
to the pilus dentilis. Remaining characters similar to Hypoaspis. 

MALE: Unknown. 



Angosomaspis multisetosus sp. n. 

FEMALE: Dorsum covered by single dorsal shield (800 (jon long and 480 (xm 
wide) covering most of the dorsum. Over a 100 minute setae (15-20 (jun) are distri- 
buted on the shield, obscuring any paired arrangement. Only four pairs of anterior 
setae and one pair of posterior marginal setae are of 'normal' Hypoaspis-type 
length and appearance (60-95 [Am). The distribution and the relative lengths of 
the setae are shown in fig. 88. The laterodorsal chaetotaxy is obscured by a row 
of setae which are inserted on the soft integument between the narrow peritrematal 
shield and the dorsal shield, in mounted specimens this part of the integument in- 
variably folds below the dorsal shield and only the examination of the mite in lateral 
position reveals that these setae are not inserted on the shield. The dorsal shield 
is finely granulated but shows no ornamentation. Tectum (fig. 91) triangulate 
with margins partly or totally denticulate. 

Tritosternum normal with long pilose laciniae. Sternal shield (190 {xm long 
and 170 fxm wide at St2) with straight anterior and posterior margin, finely granu- 
lated surface without ornamentation. Sternal setae long (e.g. St2 150 ^m) and 
whiplike, very attenuated and coming to a fine end, they are similar to the remaining 
(about 30 pairs) ventral setae. Genital shield large and elongate (distance between 
genital setae 145 (Am), devoid of ornamentation. Anal shield with slightly curved 
anterior margin, paranal setae longer than postanal seta. The peritreme extends 
anteriorly slightly beyond the middle of coxa I, it is accompanied by a very narrow 
external and internal peritrematal shield which is free both anteriorly and posteriorly. 



HYPOASPIS MITES AND BEETLES 



93 



Gnathosoma (fig. 94) with well-sclerotized corniculi and fimbriate internal 
malae with six rows of minute deutosternal denticles. Movable digit of chelicera 
(fig. 92) bidentate, fixed digit with about a dozen fine pointed denticles proximal to 
the pilus dentilis. 




92 

FIGS 88-94. Angosomaspis multisetosus gen. n., sp. n., female. Fig. 88. Dorsum. Fig. 89. 
Venter. Fig. 90. Leg IV. Fig. 91. Tectum. Fig. 92. Chelicera. Fig. 93. Tarsus 
II. Fig. 94. Gnathosoma, ventral view. 



94 M. COSTA 

The approximate lengths of the legs (excluding pretarsi) are: I 760 (xm; II 
650 (j.m; III 680 (Jim; IV 890 [xm. Long macrosetae on femora II-IV, ad\ of 
femur IV (490 [xm) extending beyond the middle of the corresponding tarsus. 
Only one macroseta (pdz) on tarsus IV (fig. 90). Tarsus II (fig. 93) with two sub- 
terminal pointed spine-like setae, usually the tip is broken off leaving a sharp 
jagged surface. 

MATERIAL: Holotype: $, ex Angosoma centaurus (Scarabaeidae : Dynastinae), 
nr. Abidjan, Ivory Coast, 1969, coll. Dr M. Remillet. Paratypes: 20 $?, same 
data as type. 

Promacrolaelaps gen. n. 

FEMALE : Large mites with a convex dorsal shield which covers the sides and the 
dorsum. Thirty-one pairs (17 podonotal and 14 opisthonotal) of long setae are 
inserted on the shield. Genital shield large, expanded posterior to the genital setae. 
Macrosetae present on femora I-IV, genua III-IV and single macroseta on tarsus IV. 
Tarsus II without spine-like or spur-like setae. Gnathosoma with seven rows of 
deutosternal denticles, fixed digit of chelicera with six medium-sized sharp teeth 
proximal to the pilus dentilis. 



MALE: Unknown. 



Promacrolaelaps hunteri sp. n. 



FEMALE: The dorsum is completely covered by a large convex dorsal shield 
(1260 |xm long and 820 [xm wide) which covers the mite also laterally. Thirty-one 
pairs of simple long setae (17 podonotal and 14 opisthonotal) are inserted on the 
shield. Setae Z4 are the longest (510 [xm) dorsal setae. The shape of the shield 
as well as the lateral displacement of many setae did not permit positional homo- 
logization of most setae, their distribution and relative lengths being shown in 
fig. 95. Tectum (fig. 97) with denticulate margin. 

Tritosternum normal, with pilose laciniae. Sternal shield (170 (xm long and 210 [xm 
wide at St2) wider than long, with long sternal setae extending considerably beyond 
the bases of consecutive setae. The shield is ornamented and granulate, its anterior 
and posterior margins are straight to slightly concave. Genital shield (fig. 96) 
large (distance between genital setae 220 (xm), well ornamented and expanding 
posterior to the genital setae. Metapodal shields small and irregular. Anal shield 
with rounded anterior margin, postanal seta slightly longer than paranal setae. 
The anal shield is flanked by a pair of long (370 [xm) setae. Approximately 17 pairs 
of setae are inserted on the integument posterior to coxae IV. The peritreme 
extends anteriorly beyond the middle of coxa I and is free both anteriorly and 
posteriorly. 

Gnathosoma (fig. 100) with well-sclerotized corniculi and fimbriate internal 
malae, seven rows of deutosternal teeth (10-20 per row) are present. Movable 
digit of the chelicera (fig. 99) bidentate, fixed digit with six well-defined, medium- 
sized sharp teeth proximal to the pilus dentilis. 



HYPOASPIS MITES AND BEETLES 



95 



The approximate lengths of the legs (excluding pretarsi) are: I 980 (xm; II 
850 fxm; III 830 (Jim; IV 930 [xm. Macrosetae present in femora I and II 
(pdi), III and IV (ad\, 450 (j.m long on femur IV) ; on genua III and IV (ad\, 450 (xm 
long on genu IV) and a single macroseta (pdz 250 yjri) on tarsus IV (fig. 101). 



'lil/'MN '' ' '' 1 ' N 

; KiVV.M v ' III > 




100 



FIGS 95-101. Promacrolaelaps hunteri gen. n., sp. n., female. Fig. 95. Dorsal shield. 
Fig. 96. Venter. Fig. 97. Tectum. Fig. 98. Tarsus II. Fig. 99. Chelicera. 
Fig. 100. Gnathosoma, ventral view. Fig. 101. Leg IV. 



96 M. COSTA 

Tarsus II (fig. 98) with simple pointed setae only, the ventral setae being slightly 
stouter. 

This species is named in honour of my friend and colleague Dr P. E. Hunter 
(Department of Entomology, The University of Georgia). 

MATERIAL: Holotype: 9 ex Promacrus bimucronatus Pallas (Scarabaeidae : 
Euchirinae), Tivon, Israel, August 20, 1967, coll. M. Costa. Paratypes: 5 $<j>, 
same data as type. 

DISCUSSION 

The vague early definition of genera and species can often be applied to several 
different taxa subsequently discovered. In the present study the early descriptions 
and figures of Hypoaspis krameri Can. apply equally well to H. krameri Can. sensu 
Evans & Till (1966), H. neokrameri sp. n. ; H. pentodoni sp. n. ; H . phyllognathi sp. n. ; 
H. integer Berlese sensu Samsinak (1960), etc. Vitzthum (1940-43) records 'H. 
krameri' from the following hosts: 0. nasicornis, Cetonia aurata, Potosia floricola, 
Pentodon punctatus, Polyphylla fullo apparently placing under H. krameri several 
species with a higher degree of host specificity than assumed by him. The un- 
certainty of the determination of mites of the Hypoaspis and Coleolaelaps com- 
plexes by Berlese has been discussed by Costa & Hunter (op. cit.). In view of this 
early species confusion and the fact that their material was obtained from a new 
host, namely Lucanus sp., it seems rather uncertain that H. krameri Can. sensu 
Evans & Till is actually conspecific with the type species, it undoubtedly agrees 
with the description and definition of Hypoaspis s. str. However, in order to avoid 
additional confusion on the subject, the decision made by Evans & Till is accepted 
here until the type (? lost) or topotypic material from the type host can be examined. 
The definition of Hypoaspis s. str. as conceived here, as well as the definition of 
Coleolaelaps Berl. as conceived by Costa & Hunter (op. cit.) will necessitate the 
transfer of many species from the latter genus to Hypoaspis s. str., this being, 
however, outside the scope of the present study. 

Being convinced that host-association is just as good a character for taxonomic 
discrimination as morphological characters, I have designated four new genera 
of which the first two (Lucanaspis and Dynastaspis) are closer related to Hypoaspis 
s. str. than the remaining two (Angosomaspis and Promacrolaelaps) . As additional 
species are expected for these genera, the generic definitions are short and not very 
rigorous, allowing amendment for the inclusion of new species. The generic 
delimitations being to a large degree a matter of personal opinion, I prefer this 
treatment to the lumping of many different forms for 'phylogenetic' reasons. 

The juvenile forms of Hypoaspis s. str. are described here for the first time (H. 
rhinocerotis and H. athasiae) and they agree to the general type of the free-living 
laelapids (vide H. aculeifer (Can.), Evans & Till, op. cit.). I should like to point 
out the 'long-haired' nature of the juveniles: the protonymph having longer setae 
than the deutonymph and this having longer setae than the adult. 

The occurrence of macrosetae on the idiosoma as well as on the legs, mainly 
leg IV, seems to have arisen independently in various mesostigmatic mites associated 



HYPOASPIS MITES AND BEETLES 97 

with arthropods. The function of the macrosetae is at present unknown, it may 
be mainly thigmotactic and concerned with finding the right location on the host, 
or it may also serve to avoid overcrowding on the host, assuring each mite its own 
volume of space. 



SUMMARY 

The following species and genera of mites associated with phytophagous lamellicorn 
beetles are described and figured: Hypoaspis neokrameri sp. n. ($); Hypoaspis 
pentodoni sp. n. ($, <) ; Hypoaspis phyllognathi sp. n. ($) ; Hypoaspis integer Berlese 
($) ; Hypoaspis rhinocerotis Ouds. ($, $, dn, pn) ; Hypoaspis athiasae sp. n. ($, <, 
dn, pn) ; Hypoaspis dubius sp. n. ($) ; Hypoaspis remilleti sp. n. (?) ; Lucanaspis 
brachypedes gen. n., sp. n. ($) ; Dynastaspis walhallae gen. n., sp. n. ($) ; Angosomaspis 
multisetosus gen. n., sp. n. (?) ; Promacrolaelaps hunteri gen. n., sp. n. ($). 

ACKNOWLEDGEMENTS 

I would like to thank the following for the loan or presentation of material used 
in this study: Dr C. Athias-Henriot (Laboratoire du Sol de I.N.R.A., Dijon); Dr 
Lee H. Herman, Jr. (American Museum of Natural History, New York); Dr M. 
Remillet (O.R.S.T.O.M., Centre d'Adiopodoume, Abidjan, Ivory Coast); Dr K. 
Samsinak (Entomological Institute of the Czechoslovak Akademy of Sciences, 
Prague) ; Dr M. J. Stelzer and Mr B. Zelazny (U.N./S.P.C. Rhinoceros Beetle Project, 
Apia, Western Samoa). I am grateful to Dr F. Pegazzano (Stazione di Entomo- 
logia Agraria, Florence) for information concerning specimens in the Berlese col- 
lection. 

It is my pleasure to express greatful thanks to Dr P. E. Hunter (Department 
of Entomology, The University of Georgia, Athens) for his generous hospitality, 
his continuous interest and for his criticism of parts of the manuscript. 

This study was made while holding a Senior Foreign Scientist Fellowship of the 
National Science Foundation at the Department of Entomology, The University 
of Georgia. 

REFERENCES 

COSTA, M. & HUNTER, P. E. (in press). The genus Coleolaelaps Berlese, 1914 (Acarina: Meso- 
stigmata) . 

EVANS, G. O. 1958. A Review of the Laelaptid Paraphages of the Myriapoda with descriptions 

of three new species (Acarina: Laelaptidae) . Parasitology 45 : 352-368. 

1963. Observations on the chaetotaxy of the legs in the free-living Gamasina (Acasi : 
Mesostigmata) . Bull. Brit. Mus. nat. Hist. (Zool.). 10 (5) : 275-303. 

EVANS, G. O. & TILL, W. M. 1966. Studies on the British Dermanyssidae (Acari: Mesostig- 
mata). Pt II Classification. Bull. Brit. Mus. nat. Hist. (Zool.). 14 (5) : 109-370. 

GRANDI, G. 1925. Contributo alia conoscenze biologica e morphologica di alcuni Lamellicorni 
fillifagi e descrizione di una nuova specie di Acaro. Boll. Lab. Zool. gen. agr. R. Scuola 
Agric. Portici 18 : 159-224. 



98 M. COSTA 

HUNTER, P. E. & COSTA, M. (in press). Gymnolaelaps shealsi n. sp. (Acarina: Mesostigmata) 

associated with the imported fire ant. 
SAMSINAK, K. 1960. Kurze Bemerkungen uber Mesostigmata (Acari). Cas. 6sl. Spol. ent. 

57 (3) : 275-284. 
SIKORA, W. B. 1968. A review of the genus Coleolaelaps Berlese, 1914 with descriptions of 

three new species (Acarina: Mesostigmata). M.S. Thesis, University of Georgia, Athens, 

Georgia, U.S.A. pp. 43. 
VITZTHUM, H. 1940-43. Acarina in H. G. Bronn (ed.), Klassen und Ordnungen des Tierreiches. 

Vol. 5, Part 5, Book 5, ion pp., Leipzig. 




Dr M. COSTA, M.Sc., Ph.D. 
KIBBUTZ MISHMAR HAEMEK 
ISRAEL 



A LIST OF SUPPLEMENTS 
TO THE ZOOLOGICAL SERIES 

OF THE BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 



1. KAY, E. ALISON. Marine Molluscs in the Cuming Collection British Museum 
(Natural History) described by William Harper Pease. Pp. 96; 14 Plates. 
1965. (Out of Print.) 3.75. 

2. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and 
Valenciennes. Pp. 180; n Plates, 15 Text-figures. 1967. 4. 

3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure of Mineralogy 
at the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125; 29 Plates, 
77 Text-figures. 1969. 4.50. 

4. HAYNES, J. R. Cardigan Bay recent Foraminifera (Cruises of the R.V. Antur) 
1962-1964. (In press.) 



Printed in England by Staples Printers Limited at their Kettering, Northants, establishment 



PI97J 

THE POLYCHAETE FAUNA OF THE 
SOLOMON ISLANDS 




P. E. GIBBS 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 21 No. 5 

LONDON : 1971 



THE POLYCHAETE FAUNA OF THE 
SOLOMON ISLANDS 




BY 

PETER EDWIN GIBBS 

Marine Biological Association of the U.K., Plymouth 



Pp. 99-211; 17 Text-figures 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 21 No. 5 

LONDON: 1971 



THE BULLETIN OF THE BRITISH MUSEUM 

(NATURAL HISTORY), instituted in 1949, is issued 
in five series corresponding to the Departments of the 
Museum, and an Historical series. 

Parts will appear at irregular intervals as they 
become ready. Volumes will contain about three or 
four hundred pages, and will not necessarily be completed 
within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 21, No. 5 of the Zoological series. 
The abbreviated titles of periodicals cited follow those 
of the World List of Scientific Periodicals. 



World List abbreviation 
Bull. Br. Mus. not. Hist. (Zool.). 



Trustees of the British Museum (Natural History), 1971 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 21 September, 1971 Price 3-55 



THE POLYCHAETE FAUNA OF THE 
SOLOMON ISLANDS 

By P. E. GIBBS 

CONTENTS 

Page 

SYNOPSIS ........... ioi 

INTRODUCTION ........ . ioi 

GENERAL ACCOUNT .......... 102 

COMPOSITION OF THE POLYCHAETE FAUNA AND ITS ZOOGEOGRAPHICAL 

AFFINITIES .......... 104 

ECOLOGICAL OBSERVATIONS 

Littoral survey . . . . . . . . . in 

Dredge survey of Marovo Lagoon . . . . . . 114 

Commensal polychaetes . . . . . . . . 116 

Brackish water and terrestrial species . . . . . 117 

Spawning of Eunice (Palola) siciliensis . . . . . 118 

SYSTEMATIC ACCOUNT . . . . . . . . . 119 

REFERENCES ........... 205 

SYNOPSIS 

An account of the polychaete fauna of the Solomon Islands (western Pacific Ocean) is given, 
based on a large collection which was obtained during the Royal Society Expedition in 1965. A 
total of 220 species are recorded. Two new genera (Family Nereidae), 13 new species and 3 new 
subspecies are described. The zoogeographical affinities of the fauna are analysed and certain 
ecological aspects discussed. 

INTRODUCTION 

As pointed out by Knox (1958), the polychaetes of the Pacific Islands are poorly 
known and much work remains to be done before a relatively complete bio- 
geographical analysis can be made. Apart from reports on small collections or 
individual families, comprehensive accounts of the polychaetes from any of the 
island groups in the tropical west Pacific regions are few. Augener (19270) records 
47 species from New Britain and Fauvel (1947) describes about 100 species from 
New Caledonia. The fauna of the Marshall Islands is comparatively well known 
with 131 species recorded as a result of the work of Hartman (1954) and, more 
recently, of Reish (1968). 

The Solomon Islands are situated between the latitudes of 5S and 12 S and 
between the longitudes of 155 E and 163 E. From the zoogeographical viewpoint 
these islands are of great interest since they lie between the faunistic centre of the 



102 P. E. GIBBS 

Indo-west-Pacific region, the Malay Archipelago, and the more distant islands of 
the Central Pacific. Thus, one of the primary aims of the Expedition organized 
by the Royal Society of London in 1965 was to determine the biogeographical 
relationships of these islands with the adjacent regions. As a member of the Marine 
Party of this Expedition the author spent five months, from July to December, 
investigating the polychaete fauna and this paper is an account of the ecological 
and faunistic observations that were made during this period. 

I am indebted to the Royal Society, Professor E. J. H. Corner, F.R.S., (Expedition 
Leader) and Professor J. E. Morton (Leader of the Marine Party) for the opportunity 
to participate in the Expedition to the Solomon Islands in 1965. For their help in 
the field I have to thank the members of the Marine Party, particularly Dr. D. R. 
Stoddart, Captain S. B. Brown and crew of the A. K. Maroro, and our native 
assistants, especially Ini Munamaori. I am also grateful to Wilson Ifunaoa for 
providing a valuable collection of the 'palolo' rising from Fanalei, Malaita. 

For their kind assistance and helpful advice in the identification of various 
specimens and families, I have to thank Dr. Olga Hartman (two new genera of 
Nereidae), Dr. Minoru Imajima (Syllidae), Dr. Charlotte Mangum (Maldanidae) and 
Professor G. P. Wells (Abarenicola) . Other groups were identified by Dr. P. 
Bergquist (Porifera), Miss A. Clark (Echinodermata) and Dr. S. J. Edmonds 
(Sipunculoidea). I am also indebted to Dr. J. D. George for the loan of specimens 
from the collections of the British Museum (Natural History) . 



GENERAL ACCOUNT 

The Marine Party of the Expedition was based and carried out investigations on 
Guadalcanal, in the Florida Islands (Nggela Group), Russell Islands and New 
Georgia Group (for a detailed account of the Expedition, see Corner, 1969). In these 
areas, collections of polychaetes were obtained from the following localities (see 
figs i and 2) : western end of Guadalcanal - Komimbo Bay, Cape Esperance and 
Naro Bay ; north coast of Guadalcanal - Mamara Point, Honiara and Lunga Point ; 
eastern end of Guadalcanal (Marau Sound) - Graham Point, Fintry Point, Marauni- 
bina Island, Pigeon Island, and Lauvie Island ; Florida Islands - Kokomtambu 
Island, Tetel (or Gaskell) Island, Nggela (or Florida) Island and Haroro; Russell 
Islands (Banika Island) - Yandina, Sifola and Lingatu ; New Georgia Group (Marovo 
Lagoon) - Matiu Island, Pirikale Island, Paleki Island and Batuona (or Wickham) 
Island. Additional collections were made at Gizo Island (New Manra) and on 
Malaita - Auki Harbour, Kalota Island, Alite Harbour and at the southern end of 
Maramasike Passage. 

The polychaete fauna of the moderate to very exposed coral reef platforms 
consists chiefly of coral-boring and crevice-dwelling species. To obtain such species 
it is necessary to break open the coral rock with hammer and chisel and then care- 
fully extract the worms from their burrows or from crevices using forceps. Those 
species living amongst the algal cover or chaetopterid tubes are best obtained by 
dissecting and then washing these materials. Investigations of these habitats were 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 



103 




FIG. i. Map of the Solomon Islands showing collection localities mentioned in the text. 
Insets - Marau Sound (left) and parts of the Russell Islands and Florida Islands (right). 



io 4 P. E. GIBBS 

made at (collection dates in brackets) Mamara Pt. (24.viii), Sifola (2.ix), Lingatu 
(19. x), Matiu and Paleki Is. (3-7.vii and 27-3i.viii), Batuona I. (3-ix) and at New 
Manra (4-5 .ix). Further collections were made at Kokomtambu I. (2.vii) and at 
Maramasike Pg. (26.xi). Beachrock fauna was investigated at Lauvie I. (iS.ix and 
4 .x). 

On the more sheltered shorelines a wide range of sedimentary deposits was studied. 
The larger species of the infauna were obtained by digging and the smaller species 
by sieving samples of the deposit through 0-5, i-o or 2-0 mm mesh, the mesh size 
depending on the grade of the sediment. Investigations of the infauna inhabiting 
a variety of deposits, ranging from thick mud to shell gravel and coral debris, were 
carried out at Tetel I., Haroro and Nggela I. (5-io.vii and 24-2g.vii), Komimbo Bay, 
Cape Esperance and Naro Bay (i8-2O.vii and i6-2i.viii), Graham Pt., Fintry Pt. 
and Maraunibina I. (i7.ix-io.x). Small collections were also made at Yandina 
(i8.x) and Honiara (28. xi). 

In addition to the above studies, further species were obtained from a brackish 
water lagoon at Lunga Point near the mouth of the Lunga River (g.ix-n.ix) and 
from encrusting organisms, particularly sponges, on wharf piles at Yandina (i9-20.x 
and 5.xi). At a number of the localities mentioned above and including Auki Hr. 
and Kalota I. (20-22. xi) commensal polychaetes were discovered. It was not 
possible for the author to be present at a rising of the 'palolo' but a small collection 
of the spawning that took place at Fanalei on 2~3.xi.66 was kindly forwarded by 
Wilson Ifunaoa. Dredging operations were confined to the New Georgia Group 
where a survey of the benthos of Marovo Lagoon was carried out during the period 
22.x-i4.ix in conjunction with the studies of Dr. D. R. Stoddart. 

The nomenclature used in this paper is that of Hartman (1959, 1965) although 
later revisions by various authors have been followed. The classification used by 
Day (1967) has been adopted. The usage of the place-names in the Solomon 
Islands is that given on the Directorate of Overseas Survey Map (i : 1,000,000). 
Other names are taken from 'the Admiralty Charts or are local names, the spelling 
of which must be regarded as provisional. 

The collections, including the type specimens, have been deposited in the British 
Museum (Natural History). 



COMPOSITION OF THE POLYCHAETE FAUNA AND ITS 
ZOOGEOGRAPHICAL AFFINITIES 

The collection of polychaetes from the Solomon Islands comprises over 2000 
specimens and is composed of 220 species.* Of this total, 88 (40%) of the species are 
represented by only one or two specimens. Thus the actual number of species in 
the fauna must be much higher, perhaps as high as 400, since probably many of the 
smaller and less conspicuous species, including many of the syllids as well as the 
paraonids and spirorbids, were overlooked in the field. The species composition of 
the fauna by families is given in Table i. The errantiate families Aphroditidae 
(24 species), Eunicidae (22) and Syllidae (20) are well represented and in total account 

* See Appendix 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 



105 



for about one-half of the errant species. Sedentary species are fewer (97 out of the 
total of 220) with the family Spionidae (15) containing the most species. In the 



TABLE i 

The species composition of the Solomon Islands fauna by families. 



Family 



No. of 
species 



Aphroditidae 24 

Palmyridae 3 

Amphinomidae 10 

Phyllodocidae 13 

Pilargidae 2 

Hesionidae 4 

Syllidae 20 

Sphaerodoridae i 

Nereidae 16 

Nephtyidae 3 

Lacydoniidae i 

Glyceridae 4 

Eunicidae 22 

Orbiniidae 2 

Spionidae 15 

Magelonidae 2 

Trochochaetidae 2 



TOTAL 



Family No. of 


species 


Chaetopteridae 
Cirratulidae 
Cossuridae 


6 

5 
i 


Flabelligeridae 
Scalibregmidae 
Opheliidae 
Sternaspidae 
Capitellidae 
Arenicolidae 


i 

2 

5 

i 

7 

i 


Maldanidae 


8 


Oweniidae 
Sabellariidae 


4 

2 


Pectinariidae 


I 


Ampharetidae 
Terebellidae 


2 
IO 


Sabellidae 


II 


Serpulidae 


9 


220 





collection, two new genera, namely Cryptonereis n.gen. and Solomononereis n.gen. 
(Family Nereidae), 13 new species and three new subspecies are present. The new 
species and subspecies are the following: 



Family Aphroditidae 

Family Hesionidae 
Family Nereidae 

Family Nephtyidae 
Family Eunicidae 
Family Spionidae 



Family Trochochaetidae 
Family Flabelligeridae 
Family Opheliidae 
Family Oweniidae 
Family Terebellidae 



Hololepidella ophiuricola n.sp. 
Lepidasthenia guadalcanalis n.sp. 
Gyptis maraunibinae n.sp. 
Cryptonereis malaitae n.gen., n.sp. 
Solomononereis marauensis n.gen., n.sp. 
Nephtys (Aglaophamus) munamaorii n.sp. 
Eunice marovoi n.sp. 
Dispio maroroi n.sp. 
Polydorella novaegeorgiae n.sp. 
Prionospio tetelensis n.sp. 
Scolelepis squamata mendanai n. subsp. 
Poecilochaetus serpens honiarae n. subsp. 
Diplocirrus glaucus orientalis n. subsp. 
Ophelia koloana n.sp. 
Myriochele heruensis n.sp. 
Pista dibranchis n.sp. 



io6 



P. E. GIBBS 



The species collected in the Solomon Islands are listed in Table 2. From the 
material three species - all maldanids - can be identified to the subfamily level only, 
17 are referable to their genus and a further six can be given near identifications to 
comparable species. Excluding these species and also the new species and sub- 
species a total of 178 species is available for analysis. 



TABLE 2 

List of species taken in the Solomon Islands. 



APHRODITIDAE 

Gastrolepidia clavigera Schmarda 

Harmothoe nigricans Horst 

Harmothoe sp. 

Hololepidella nigropunctata (Horst) 

H . ophiuricola n.sp. 

Iphione muricata (Savigny) 

Lepidasthenia elegans (Grube) 

L. maculata Potts 

L. guadalcanalis n.sp. 

L. microlepis Potts 

L. stylolepis Willey 

Lepidonotus (Thormora) jukesi (Baird) 

Paradyte crinoidicola (Potts) 

Paralepidonotus ampulliferus (Grube) 

P. indicus (Potts) 

Polyodontes maxillosus (Ranzani) 

P. melanonotus (Grube) 

Psammolyce zeylanica Willey 

Sigalion bandaensis Horst 

Sthenelais heterochela Horst 

S. zeylanica Willey 

Sthenelanella ehlersi (Horst) 

Sthenolepis japonica (Mclntosh) 

Thalenessa digitata Mclntosh 

PALMYRIDAE 

Bhawania goodei Webster 
B. pottsiana Horst 
Paleanotus debilis (Grube) 

AMPHINOMIDAE 

Amphinome nigrobranchiata Horst 

Chloeia conspicua Horst 

Euphrosine foliosa Aud. & M.-Ed. 

E. myrtosa Savigny 

Eurythoe complanata (Pallas) 

? Eurythoe sp. 

Notopygos gregoryi Holly 

N. sibogae Horst 

Pareurythoe pitipanaensis De Silva 

Pseudeurytlwe paucibranchiata Fauvel 



PHYLLODOCIDAE 

Eteone japanensis Mclntosh 

Eulalia albopicta Marenzeller 

E. viridis (Linnaeus) 

E. (Pterocirrus) magalhaensis Kinberg 

Notophyllum splendens (Schmarda) 

Phyllodoce fristedti Bergstrom 

P. malmgreni Gravier 

P. pruvoti Fauvel 

P. quadraticeps Grube 

P. (Anaitides) madeirensis Langerhans 

P. (Anaitides) parva Hartmann-SchrSder 

P. (Genetyllis) castanea (Marenzeller) 

P. (Genetyllis) gracilis Kinberg 

PILARGIDAE 

Sigambra hanaokai (Kitamori) 
Synelmis albini (Langerhans) 

HESIONIDAE 

Gyptis capensis (Day) 
G. maraunibinae n.sp. 
Hesione splendida Savigny 
Leocrates chinensis Kinberg 

SYLLIDAE 

Autolytus sp. 

Brania clavata (Claparede) 
Exogone gemmifera Pagenstecher 
E. uniformis Hartman 
E. verugera (Claparede) 
Haplosyllis spongicola (Grube) 
Langerhansia cornuta (Rathke) 
L.? rosea (Langerhans) 
Sphaerosyllis hirsuta Ehlers 
Syllis longissima Gravier 
Syllis sp. cf. gracilis Grube 
Trypanosyllis coeliaca Claparede 
T. (Trypanedenta) sp. 
Typosyllis alternata (Moore) 
T. armillaris (Miiller) 
T. brachycola (Ehlers) 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 



107 



T. exilis (Gravier) 
T. lucida (Chamberlin) 
T. prolifera (Krohn) 
Ty posy His sp. 

SPHAERODORIDAE 

Sphaerodoridium claparedii (Greeff) 

NEREIDAE 

Ceratonereis costae (Grube) 

C. erythraeensis Fauvel 

C. mirabilis Kinberg 

Cryptonereis malaitae n.g.n.sp. 

Namalycastis indica (Southern) 

Nereis (Neanthes} caudata (Delle Chiaje) 

N. (Neanthes) unifasciata Willey 

N. (Neanthes} cf. kerguelensis Mclntosh 

Perinereis cultrifera (Grube) 

P. nigropunctata (Horst) 

P. nuntia (Savigny) 

Platynereis insolita Gravier 

Pseudonereis anomala Gravier 

P. masalacensis (Grube) 

P. variegata (Grube) 

Solomononereis marauensis n.g.n.sp. 

NEPHTYIDAE 

Nephtys (Aglaophamus) munamaorii n.sp. 
N. (Micronephthys} sphaerocirrataWesenberg- 

Lund 
N. (Nephtys) sp. cf. palatii Gravier 

LACYDONIIDAE 

Paralacydonia weberi Horst 

GLYCERIDAE 

Glycera gigantea Quatrefages 
G. lancadivae Schmarda 
G. longipinnis Grube 
G. rouxi Aud. & M.-Ed. 

EUNICIDAE 

Arabella tricolor (Montagu) 

A. mutans (Chamberlin) 

Dorvillea sp. 

Drilonereis major Crossland 

Eunice afra Peters 

E. antennata (Savigny) 

E. aphroditois (Pallas) 

E. coccinea Grube 

E. grubei Gravier 

E. marovoi n.sp. 

E. norvegica (Linnaeus) 



E. tentaculata Quatrefages 

E. tubifex Crossland 

E. (Palola) siciliensis Grube 

Lumbrineris latreilli Aud. & M.-Ed. 

L. papillifera (Fauvel) 

L. sphaerocephala (Schmarda) 

Lysidice collaris Grube 

Marphysa macinloshi Crossland 

Nematonereis unicornis (Grube) 

Oenone fulgida (Savigny) 

Onuphis (Nothria) holobranchiata Marenzeller 

ORBINIIDAE 

Haploscoloplos bifurcatus Hartman 
Naineris laevigata (Grube) 

SPIONIDAE 

Dispio maroroi n.sp. 

Laonice cirrata (Sars) 

Malacoceros indicus (Fauvel) 

Nerinides sp. cf. gilchristi Day 

Polydorella novaegeorgiae n.sp. 

Prionospio cirri/era Wir6n 

P. ehlersi Fauvel 

P. malmgreni Claparede 

P. pinnata Ehlers 

P. steenstrupi malayensis Caullery 

P. tetelensis n.sp. 

Pseudopolydora corallicola Woodwick 

Pseudopolydora sp. 

Scolelepis squamata mendanai n.subsp. 

Spio filicornis (Miiller) 

MAGELONIDAE 

Magelona japonica Okuda 
Magelona sp. 

TROCHOCHAE TIDAE 

Poecilochaetus serpens honiarae n.subsp. 
P. tropicus Okuda 

CHAE TOP TERIDAE 

Chaetopterus variopedatus (Renier) 
Mesochaetopterus Sagittarius (Claparede) 
Phyllochaetopterus elioti Crossland 
P. herdmani (Hornell) 
P. socialis Claparede 
Spiochaetopterus costarum costarum 
(Claparede) 

CIRRATULIDAE 

Cirriformia filigera (Delle Chiaje) 
C. punctata (Grube) 



io8 



P. E. GIBBS 



Dodecaceria fistulicola Ehlers 
D. laddi Hartman 
Tharyx sp. 

COSSURIDAE 

Cossura coasta Kitamori 

FLABELLIGERIDAE 

Diplocirrus glaucus orientalis n.subsp. 

SCALIBREGMIDAE 

Hyboscolex longiseta Schmarda 
Scalibregma inflatum Rathke 

OPHELIIDAE 

Armandia lanceolata Willey 
A. leptocirrus (Grube) 
A. longicaudata (Caullery) 
Ophelia koloana n.sp. 
Polyophthalmus pictus (Dujardin) 

STERN ASPIDAE 

Sternaspis scutata (Renier) 

CAPITELLIDAE 

Capitellethus dispar (Ehlers) 
Capitobranchus sp. 
Dasybranchus caducus (Grube) 
Mastobranchus dollfusi Fauvel 
M. trinchesii Eisig 
Mediomastus sp. cf. capensis Day 
Notomastus sp. 

ARENICOLIDAE 

Abarenicola claparedii claparedii (Levinsen) 

MALDANIDAE 

Clymenella (= Macroclymene) sp. i 
Clymenella (= Macroclymene) sp. 2 
Clymenella (= Euclymene) sp. 
Euclymeninae sp. A. 
Euclymeninae sp. B. 
Euclymeninae sp. C. 
? Nicomache sp. 
Praxillella sp. 

OWENIIDAE 

Myriochele eurystoma Caullery 
M. heruensis n.sp. 



Myriochele sp. 

Owenia fusiformis Delle Chiaje 

SABELLARIIDAE 

Lygdamis ehlersi (Caullery) 
L. indicus Kinberg 

PEC TINARHDAE 

Pectinaria (Pectinaria) antipoda Schmarda 

AMPHA RE TIDAE 

Isolda pulchella Miiller 
? Sosane wireni Caullery 

TEREBELLIDAE 

Amaeana trilobata (Sars) 
Euthelepus kinsemboensis Augener 
Eupolymnia nebulosa (Montagu) 
Loimia medusa (Savigny) 
Lysilla ubianensis Caullery 
Pista dibranchis n.sp. 
P. typha (Grube) 

Reteterebella queenslandia Hartman 
Terebella ehrenbergi Grube 
Terebellides stroemi Sars 

SABELLIDAE 

Branchiomma cingulata (Grube) 

Hypsicomus phaeotaenia (Schmarda) 

Megalomma intermedium (Beddard) 

M. linaresi (Rioja) 

M. quadrioculatum (Willey) 

M. trioculatum Reish 

M. vesiculosum (Montagu) 

Potamilla ehlersi Gravier 

Sabella fusca Grube 

5. melanostigma Schmarda 

Sabellastarte sanctijosephi (Gravier) 

SERPULIDAE 

Filograna implexa Berkeley 
Hydroides minax (Grube) 
H. uncinata (Philippi) 
Mercierella enigmatica Fauvel 
Serpula hartmanae Reish 
S. vermicularis Linnaeus 
Spirobranchus coutierei (Gravier) 
5. giganteus (Pallas) 
Vermiliopsis glandigerus Gravier 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 



109 



At this stage a zoogeographical analysis of the Solomon Islands fauna can be 
regarded only as provisional since many areas within the Indo-Pacific still remain 
to be investigated and further records, as well as taxonomic revisions, will modify 
the known distribution patterns of many species. The data for this analysis are 
given in Table 3. The faunistic regions outlined in this table are necessarily broad: 
in the Pacific region, only those island groups for which the polychaete faunas are 
fairly well known are considered. As far as possible, synonymies have been taken 
into account. In compiling the distributional data for each species the following 
references have been consulted : 



(i) Eastern Atlantic 

(ii) Mediterranean Sea 
(iii) South Africa 
(iv) East Africa (Red Sea to 

32S) 

(v) Indian waters (Persian 
Gulf to Andaman Is.) 

(vi) South-west Australia 

(vii) Malay Archipelago (Malaya 
to New Guinea and 
including Philippine and 
Palau Is.) 

(viii) East Australia (south 
to Port Jackson) 

(ix) New Caledonia 
(x) Central Pacific (Society 
and adjacent Islands) 

(xi) New Zealand 

(xii) Marshall Islands 
(xiii) Hawaii 
(xiv) Japan 



Augener, 1918; Fauvel, 1923, 1927; Kirke- 

gaard, 1959. 
Fauvel, 1923, 1927. 
Day, 1967. 
Crossland, 1903, 1904, 1924; Day, 1962, 1967; 

Gravier 1900-1908. 
De Silva, 1961, 1965, 19650; Fauvel, 1953; 

Tampi & Rangarajan, 1964; Wesenberg- 

Lund, 1949. 
Augener, 1913, 1914; Fauvel, 1922; Kott, 

1949. 
Caullery, 1944; Ehlers, 1920; Fauvel, 1935, 

1939; Horst, 1910-1924; Mesnil & Fauvel, 

1939; Okuda 19370; Pillai, 1965. 

Augener, 1927; Monro, 1931; Rullier, 1965; 

Russell, 1962; Straughan, 1967. 
Fauvel, 1947. 
Fauvel, 1947; Monro, 1928, 1939, 19390. 

Augener, 1924, 1926; Ehlers, 1904; Knox, 

1951, 19510, 1960. 
Hartman, 1954; Reish, 1968. 
Hartman, 1966; Straughan, 1969. 
Imajima & Hartman, 1964; Imajima, 1966- 

1967. 



Of the total of 178 species recorded from the Solomon Islands, 21 or n-8% are 
considered to be cosmopolitan in their distribution and can be eliminated from the 
analysis in order to obtain a clearer pattern of the faunal relationships. 

As expected, the Solomon Islands fauna has a large overlap with the faunas of the 
Malay Archipelago (61-1%) and of Indian waters (64-3%), and over one-half (55-4%) 
of the species are known to extend to the east coast of Africa. Thus the Indo-west- 
Pacific component forms a high proportion of the Solomon Islands fauna. In the 
Pacific region, it is interesting that the number of species in common between the 



P. E. GIBBS 



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THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS in 

Solomon Islands and other Island groups lying within the tropics decreases with 
increasing distance: the faunas of both New Caledonia and the Marshall Islands 
show similar affinities (27-4% and 29-3% respectively) as do the faunas of the 
islands of the Central Pacific (15-9%) and Hawaii (15-9%). A relatively high 
proportion of the Solomon Islands species extend to Japan (31-2%) but fewer than 
expected have been recorded from the east coast of Australia (26-1%) although this 
percentage should probably be much higher. 

It is difficult to compare these results with those of previous analyses because of 
the lack of earlier records from the tropical west Pacific. For example, Knox (1958) 
had records of only 81 species from this region (Micronesia - Melanesia) for his analysis 
of the distribution of polychaetes in the Indo-Pacific region and of this total 27% 
were Indo-Pacific and 31% were cosmopolitan. However the larger number of 
records from the present survey indicates that the Indo-Pacific element is proportion- 
ally much greater. Interestingly, previous estimates of the Indo-Pacific element 
in the faunas of New Zealand and Japan, of 12 % and 30 % respectively (Knox, 1963 ; 
Imajima & Hartman, 1964), closely correspond to the overlaps calculated above 
between these two regions and the Solomon Islands, the fauna of the latter being 
essentially Indo-Pacific. 

The records from the Solomon Islands have extended the known distribution of 
thirteen species from the Indian Ocean to the West Pacific region. These species 
are Lepidasthenia stylolepis, Psammolyce zeylanica, Pareurythoe pitipanaensis, 
Phyllodoce fristedti, Syllis longissima, Platynereis insolita, Lumbrineris papillifera, 
Marphysa macintoshi, Prionospio ehlersi, Phyllochaetopterus elioti, P. herdmani, 
Spiochaetopterus costarum costarum and Megalomma quadrioculatum. A further six 
species were known only from Atlantic or Mediterranean waters, namely Trypano- 
syllis coeliaca, Sphaerodoridium claparedii, Mastobranchus dollfusi, M. trinchesii, 
Abarenicola claparedii, and Megalomma linaresi. 

As noted above the fauna of the Solomon Islands appears to be essentially Indo- 
Pacific and apart from the new species and subspecies that were discovered, relatively 
few of the species remain unrecorded from the Indian Ocean. This Pacific element 
in the fauna is represented by such species as Notopygos gregoryi, Eteone japanensis, 
Phyllodoce pruvoti, P. (Anaitides) parva, Sigambra hanaokai, Exogone uniformis, 
Typosyllis brachycola, T. lucida, Pseudonereis masalacensis, Haploscoloplos bifurcatus, 
Pseudopolydora corallicola, Poecilochaetus tropicus, Reteterebella queenslandia, 
Megalomma trioculatum and Serpula hartmanae. However most of these species 
have few records and probably have yet to be discovered elsewhere. 

ECOLOGICAL OBSERVATIONS 

Littoral survey 

Although it is not practicable to list fully the species taken at each of the littoral 
stations, the short accounts that follow are presented as an outline of the ecological 
assemblages of species encountered in several of the more important habitats, 
in particular, those found on the reef platforms along the more exposed shorelines 



H2 P. E. GIBBS 

and those composing the infauna of certain sedimentary deposits present in 
sheltered coastal areas. The habitats of each species are given in the Systematic 
Account. The tidal range in the Solomon Islands is small, being about i-om in 
amplitude at springs, and investigations were generally concentrated betwen mid-tide 
level and low water mark. 

The importance of the boring activities of polychaetes was recognized by Gardiner 
(1903) who regarded them as the "prime and most effective agents" in the destruction 
of coralline rocks. This view has been supported by Johnson (in Hartman, 1954). 
Boring is effected chiefly by the abrasive action of either hard pharyngeal structures, 
such as possessed by the nereids and eunicids, or by modified setae, as in certain 
spionids and cirratulids (Hartman, 1954). However as pointed out by Utinomi 
(1953) it is often difficult to distinguish between those species which actually bore 
into the coral from those which utilize and perhaps enlarge the vacated burrows made 
by boring species. For example, from its habit the sabellid Hypsicomus phaeotaenia 
appears to be a boring species but it does not possess hard chitinized structures that 
could be used for penetration. Whether chemical action is employed by such a 
species has yet to be investigated. Few polychaetes contribute to reef formation 
by virtue of their construction of massed tubes and in the Solomon Islands only 
Spirobranchus giganteus appears to be sufficiently common to be considered a minor 
contributor to reef growth. 

On the reef platforms at moderately or very exposed localities, such as at Matiu, 
Batuona and Gizo, sedentary species are surprisingly few although quite extensive 
colonies of Phyllochaetopterus socialis are often present in places. On the other hand, 
errant species are fairly numerous, particularly the nereids and eunicids, which form 
the bulk of the destructive species, including Perinereis nigropunctata, Pseudonereis 
anomala, P. masalacensis, P. variegata and perhaps Platynereis insolita, together 
with Eunice afra, E. antennata, E. coccinea, E, teniaculata, E. (Palola) siciliensis 
and Lysidice collaris. Other species which may contribute to reef breakdown 
are Nematonereis unicornis, Arabella mutans, Dodecaceria fistulicola and Hypsicomus 
phaeotaenia. Many of these boring species are also common amongst the surface 
cover on the reef, that is, amongst algal growth (e.g. Amphiroa), or Phyllochaetopterus 
tubes, where they accompany the crevice-dwelling species which utilize such shelter 
when the reef is exposed at low-tide. The crevice-dwelling species are many and are 
represented by Lepidonotus (Thormora) jukesi, Paleanotus debilis, Eurythoe complanata, 
Euphrosine myrtosa, Eulalia (Pterocirrus) magalhaensis, Phyllodoce fristedti, P. pruvoti, 
P. quadraticeps , Synelmis albini, Exogone gemmifera, Syllis longissima, Trypanosyllis 
coeliaca, Typosyllis alternata, T. armillaris, T. brachycola, T. prolifera, Cirriformia 
punctata and Terebella ehrenbergi. Encrusting forms include V ermiliopsis glandigerus 
and Hydroides minax. An inconspicuous species of this habitat is Bhawania goodei 
which, although found free-living, most frequently occurs as a commensal in the 
burrows of the sipunculids Aspidosiphon elegans, Cloeosiphon aspergillum and 
Phascolosoma albolineatum. 

In sheltered localities the reef platforms are generally covered by a layer of 
sediment which is often composed of a wide variety of deposit grades ranging from 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 113 

sticky mud to coarse coral debris. At Tetel Island it was possible to investigate an 
assortment of deposits in a small area and the following lists of species discovered in 
four different grades of sediment give an indication of the diversity of the polychaete 
infauna. 

(i) In muddy silt-sand at about mid-tide level: iphione muricata, Lepidasthenia 
elegans, L. maculata, Sthenelais zeylanica, Eurythoe complanata, Eteone japanensis, 
Phyllodoce malmgreni, Langerhansia cornuta, Onuphis (Nothria) holobranchiata, 
Malacoceros indicus, Poecilochaetus serpens honiarae n. subsp., P. tropicus, 
Mesochaetopterus Sagittarius, Phyllochaetopterus elioti, P. herdmani, Pista dibranchis 
n.sp., P. typha and Megalomma vesiculosum. L. elegans and L. maculata were dis- 
covered cohabiting the tubes of M . Sagittarius and P. herdmani respectively. Living 
under coral boulders lying on this fine deposit, further species were uncovered, 
namely, Pseudeurythoe paucibranchiata, Typosyllis alternata, T. brachycola, Perinereis 
cultrifera, P. nuntia, Glycera lancadivae, Nematonereis unicornis, Arabella iricolor, 
Spiochaetopterus costarum costarum, Hyboscolex longiseta, Dasybranchus caducus, 
Lysilla ubianensis and Loimia medusa. 

(ii) In coarse coral debris, chiefly Acropora fragments, towards low water mark: 
Eurythoe complanata, Notopygos sibogae, Pseudeurythoe paucibranchiata, Leocrates 
chinensis, Ceratonereis mirabilis, Eunice aphroditois, E. (Palola) siciliensis, Cirriformia 
filigera, Armandia lanceolata, Loima medusa, Reteterebella queenslandia, Branchiomma 
cingulata, Hypsicomus phaeotaenia, Megalomma intermedium, M. linaresi, Sabellastarte 
sanctijosephi, Spirobranchus giganteus and S. coutierei. 

(iii) In shell gravel at low water mark: Lepidonotus (Thormora) jukesi, Amphinome 
nigrobranchiata, Pareurythoe pitipanaensis, Leocrates chinensis, Glycera lancadivae, 
Eunice (Palola) siciliensis, Onuphis (Nothria) holobranchiata, Mesochaetopterus 
sasittarius and Dasybranchus caducus. 

(iv) In coarse coral sand just below low water mark: Eurythoe complanata, 
Magelona japonica, Poecilochaetus serpens honiarae n. subsp., P. tropicus, Meso- 
chaetopterus Sagittarius, Spiochaetopterus costarum costarum, Capitobranchus sp. 
Dasybranchus caducus, Mastobranchus trinchesii, PNicomache sp., Pista dibranchis 
n. sp. and Megalomma vesiculosum. 

At Graham Point in Marau Sound (east Guadalcanal) extensive deposits of silty 
sand mixed with shell gravel are colonised by marine angiosperms (Thalassia and 
others) and support a rich and varied fauna. Of the polychaetes encountered here 
the most interesting are three Lepidasthenia species which were found living as 
commensals. L. microlepis and L. stylolepis were both discovered in the burrows 
of the sipunculid Siphonosoma vastus while the third species, L. guadalcanalis n.sp., 
was an inhabitant of the burrows of a large enteropneust (probably Balanoglossus 
carnosus). The polychaete infauna comprises the following species: Polyodontes 
maxillosus, Psammolyce zeylanica, Sthenelais zeylanica, Amphinome nigrobranchiata, 
Eurythoe complanata, Pseudeurythoe paucibranchiata, Glycera gigantea, G. lancadivae, 
Eunice (Palola} siciliensis, Naineris laevigata, Dispio maroroi n.sp., Scolelepis 
squamata mendanai n.subsp., Poecilochaetus tropicus, Mesochaetopterus Sagittarius, 
Dasybranchus caducus, Mastobranchus trinchesii, Clymenella (Macroclymene) sp. and 



Ii 4 P. E. GIBBS 

Pista dibranchis n.sp. Many of these species are also found below coral boulders 
together with further species including iphione muricata, Sigalion bandaensis, 
Notopygos sibogae, Pareurythoe pitipanaensis, Phyllodoce madeirensis, Notophyllum 
splendens, Synelmis albini, Gyptis maraunibinae n.sp., Hesione splendida, Ty posy His 
brachycola, Perinereis nigropunctata, Eunice antennata, E. grubei, Lysidice collaris, 
Lumbrineris latreilli, Eupolymnia nebulosa, and Euthelepus kinsemboensis. In this 
habitat, Hololepidella nigropunctata and H. ophiuricola n.sp. are also found but 
living as commensals on ophiuroids. 

Dredge Survey of Marovo Lagoon 

Along the north-east coast of New Georgia Island and around the north and east 
coasts of Vangunu Island in the New Georgia Group, elevated barrier reefs form the 



\Tokavai Lagoon 
Grassi Lagoon 




FIG. 2. Collection localities and positions of the dredge stations in 
Marovo Lagoon, New Georgia Group. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 115 

seaward edge of a more or less continuous lagoon. This lagoon is about 10 km across 
at its widest point, and its depth is less than 50 m over most of its area. Marovo 
Lagoon forms the central part of this lagoon, the northern extension being Tokavai 
and Grassi Lagoons and the southern part Kolo Lagoon. However, for convenience, 
these lagoons will be collectively referred to as Marovo Lagoon. The geomorphology 
of the region has been described by Stoddart (1969). 

A benthic survey of Marovo Lagoon was carried out in October and November 
1965. Qualitative samples of the bottom deposits were taken at 40 stations in 
depths from 2 to 35 m, using a small naturalist's dredge. Two stations (ML 218 and 
283) were worked in the northern part of the lagoon, the rest being situated in the 
central and eastern parts (fig. 2) . The fauna was extracted from the samples (between 
10 and 20 1 of sediment) by sieving through a mesh of i-o or 2-0 mm diameter. The 
bottom deposits vary from thick terrigenous mud, which occurs close to the main- 
land, to coarse coralline sands which are found along the inner margins of the outer 
reefs. The station data are given in Table 4. 



TABLE 4 
Dredge survey of Marovo Lagoon : station data. See fig. 2 for station positions. 



Station 


Date 


Depth 


Deposit 


(ML) 


(1965) 


(m) 




29 


22.X. 


22 


Silty mud 


37 


23-x. 


22 


Mud 


38 


23.X. 


9 


Silty mud 


39 


25-x. 


n 


Mud 


40 


25.X. 


18 


Mud 


4i 


25-x. 


20 


Mud 


55 


26.X. 


n 


Silty mud 


56 


26.X. 


9 


Silty mud 


68 


27. x. 


2 


Mud 


69 


27. x. 


13 


Silty mud 


72 


27. x. 


35 


Silty sand 


96 


28.X. 


9 


Sand 


97 


28.X. 


9 


Sand 


98 


28.X. 


II 


Sand 


100 


28.X. 


33 


Mud 


no 


28.X. 


26 


Mud 


116 


29.X. 


15 


Silty sand 


117 


29.X. 


5 


Sand 


ix8 


29.X. 


2 


Sand 


i34 


29.X. 


16 


Foram sand 



Station 


Date 


Depth 


Deposit 


(ML) 


(1965) 


(m) 




155 


30.x. 


9 


Mud 


156 


30-x. 


18 


Mud 


157 


3O.x. 


13 


Mud 


1 88 


i.xi 


18 


Mud 


190 


i.xi 


5 


Sand 


191 


i.xi 


2 


Silty sand 


192 


i.xi 


18 


Coarse sand 


194 


2.xi 


n 


Mud 


195 


2.X1 


n 


Mud 


196 


2.xi 


2 


Silty sand 


203 


3-xi 


4 


Coarse sand 


204 


3-xi 


18 


Coarse sand 


218 


8.xi 


18 


Mud 


228 


lo.xi 


5 


Mud 


229 


10. xi 


5 


Sand 


230 


lo.xi 


24 


Silty sand 


283 


1 3-xi 


2 


Coarse sand 


294 


14. xi 


4 


Coarse sand 


295 


14. xi 


4 


Coarse sand 


296 


14. xi 


2 


Coarse sand 



A total of 65 species were taken during the survey of which 43 were not recorded 
from the littoral zone. This category includes all the species records of the families 
Nephtyidae (3 species), Lacydoniidae (i), Cossuridae (i), Flabelligeridae (i), Sternas- 
pidae (i), Oweniidae (4), Pectinariidae (i), and Ampharetidae (2). In addition the 



u6 P. E. GIBBS 

survey provided specimens of four new species, namely, Nephtys (Aglaophamus) 
munamaorii n.sp., Polydorella novaegeorgiae n.sp., Ophelia koloana n.sp. and 
Myriochele heruensis n.sp. and also one new subspecies, Diplocirrus glaucus orientalis. 
Of the species taken, ten have not been recorded from the West Pacific, outside of the 
Malay Archipelago (Sthenelais heterochela, S. zeylanica, Sthenelanella ehlersi, Chloeia 
conspicua, Prionospio steenstrupi malayensis, Armandia longicaudata, Myriochele 
eurystoma, Isoldapulchella, Sosane wireni and Lysilla ubianensis) and the distributions 
of two species (Lumbrineris papillifera and Prionospio ehlersi} are extended from the 
coast of East Africa. A further two species, namely, Pseudopolydora corallicola and 
Megalomma trioculatum, have been described only recently from the Marshall Islands. 
Haploscoloplos bifurcatus was previously known only from south and south-east 
Australia. 

The survey provides some indications of the distribution of the commoner species 
with respect to the bottom deposits. Arbitrarily defining a "common" species as 
one that was present at three or more stations, the following appear to be mainly 
restricted to the finer grades of sediments (mud and fine sands) Sthenelais heterochela, 
Sthenolepis japonica, Pseudeurythoe paucibranchiata, Sigambra hanaokai, Nephtys 
(Aglaophamus) munamaorii n.sp., Paralacydonia weberi, Haploscoloplos bifurcatus, 
Prionospio ehlersi, P. pinnata, P. tetelensis n.sp., Diplocirrus glaucus orientalis n. 
subsp., Armandia lanceolata, Mastobranchus dollfusi, Isoldapulchella, Loimia medusa, 
and Terebellides stroemi. The coarser deposits (medium to coarse sands) appear to 
support sparse polychaete populations and only Owenia fusiformis was taken in any 
numbers. Species that tolerate a wide range of deposit grades include Glycera 
gigantea, G. lancadivae, Eunice marovoi n.sp., Prionospio steenstrupi malayensis, 
Armandia leptocirrus, Pectinaria (Pectinaria) antipoda, and Pista dibranchis n.sp. 
Apart from these common species, several appear to be locally abundant, to judge 
from the large numbers present at single stations. For example, Onuphis (Nothria) 
holobranchiata was abundant in the sample of the thick mud at ML 69. Small 
species such as Polydorella n'ovaegeorgiae n.sp. and Myriochele heruensis n.sp. were 
probably much more abundant than the sample figures at ML 39 and ML 134 suggest, 
no doubt many specimens being lost during sieving. 



Commensal polychaetes 

During the routine sampling of a number of habitats, a total of twelve species, 
mainly polynoids, were discovered living in association, apparently as commensals, 
with other animals, principally echinoderms and sipunculids. Observations on these 
associations have been presented in an earlier paper (Gibbs, 1969) but for the sake 
of completeness the commensal species and their hosts are summarized in Table 5. 

Although Macnae & Kalk (1962) have described similar associations from the 
coast of Mosambique, probably many have yet to be discovered throughout the Indo- 
Pacific region. Undoubtedly the echinoderms which act as hosts for G. clavigera, 
Hololepidella spp. and P. crinoidicola, are more numerous than the present records 
indicate. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 117 

TABLE 5 

Summary of the commensal polychaetes and their hosts in the Solomon Islands. Abbreviations : 
Po - Polychaeta ; Si - Sipunculoidea ; Ga - Gastropoda ; Oph - Ophiuroidea ; Ho - Holothuroidea; 

Cr - Crinoidea. 
Commensal Host(s) 

Bhawania goodei Aspidosiphon elegans (Si) 

Cloeosiphon aspergillum (Si) 
Phascolosoma albolineatum (Si) 

B. pottsiana Eurythoe complanata (Po) 

Eunice marovoi n.sp. Cerithium vertagus (Ga) 

Gastrolepidia clavigera Bohadschia argus (Ho) 

B. graffei (Ho) 

Holothuria (Halodeima) atra (Ho) 
Stichopus chloronotus (Ho) 
Thelenota ananas (Ho) 

Hololepidella nigropunctata Ophiarthrum elegans (Oph) 

Ophiocoma brevipes (Oph) 
O. insularia forma dentata (Oph) 
H. ophiuricola n.sp. Macrophiothrix koehleri (Oph) 

Ophiarthrum pictum (Oph) 

Lepidasthenia elegans Mesochaetopterus Sagittarius (Po) 

L. guadalcanalis n.sp. Enteropneust (Balanoglossus carnosus?) 

L. maculata Phyllochaetopterus herdmani (Po) 

L. microlepis Paraspidosiphon cumingi (Si) 

Siphonosoma vastus (Si) 

L. stylolepis Siphonosoma vastus (Si) 

Paradyte crinoidicola Himerometra robustipinna (Cr) 

Recently, Dr R. U. Gooding kindly sent the author three specimens of H. nigropunctata taken from two 
A canthaster planci L. collected at Kira-Kira, San Cristobal I. (3.xii.6g) and a further specimen taken from 
Diadema savignyi Michelin collected on the Reef Is., Santa Cruz Is. (8.xii.6g). 

Brackish water and terrestrial species 

Investigations of brackish water were mainly confined to a lagoon at Lunga 
Point, near the mouth of the Lunga River on the north coast of Guadalcanal. At 
the time of examination (g.ix-n.ix) the salinity of the lagoon was 5'6% , whilst that 
of the outside seawater was 34'5% . Three polychaete species were discovered in this 
habitat, namely Namalycastis indica, Pseudopolydora sp. and Mercierella enigmatica. 
Specimens of N. indica were found burrowing amongst the fibres composing the 
outer husk of the Nipa palm nut, many of which were lying waterlogged along the 
water's edge. The two other species were present in large numbers within the 
interstices of a sponge that was growing around submerged tree roots. The specimens 
of Pseudopolydora have proved to represent a new species (W. J. Light, personal 
communication) and those of the cosmopolitan M. enigmatica are also of interest 
in that they show the characters of Neopomatm and thus support Straughan's (1966) 
evidence for the belief that Neopomatus is the warm water form of Mercierella. 
Further details of these two species are given below in the Systematic Account. 

Wesenberg-Lund (1958) has shown that the majority of freshwater polychaetes 



n8 P. E. GIBBS 

are nereids and thus it is not surprising that those species which have been discovered 
in damp terrestrial habitats also belong to this family, in particular to the subfamily 
Namanereinae. In view of the fact that terrestrial forms are well documented from 
the East Indies (see Feuerborn, 1932 ; Lieber, 1931 ; Pflugfelder, 1933) their discovery 
in the nearby Solomon Islands is to be expected. 

Two small specimens belonging to the Namanereinae were separately collected by 
members of the Expedition Land Party (Dr. J. Greenslade and Mr. J. Peake) on 
Mt. Austen, Guadalcanal and on Kolombangara in the New Georgia Group. Both 
were taken from moist leaf litter at an altitude of 350 m. Unfortunately the identity 
of these important specimens is uncertain due to damage and although they closely 
resemble Namanereis amboinensis (Pflugfelder) they may belong to Cryptonereis 
malaitae n.gen., n.sp., described below, which may also be a damp terrestrial form. 
Until further specimens from relatively high altitudes are available the status of 
these specimens must remain undetermined. Any information concerning their 
mode of reproduction would be of great interest. 



Spawning of Eunice (Palola) siciliensis 

The spawning of the circumtropical species Eunice (Palola} siciliensis is well 
documented for the Pacific region and accounts of this phenomenon, which usually 
takes place in October or November, have been given by many authors, notably 
Burrows (1945, 1955), Gaspers (1961), Gravier (1924), Miller & Pen (1959), Stair (1847) 
and Woodworth (1907). Although the Samoan name 'palolo' is the most commonly 
used for this species, it has a variety of names, being known as 'paroro' on Rotuma, 
'balolo' in the Fiji Islands, and 'hundu' in the New Hebrides. In the Solomon Islands 
it has several names ; in the Florida Islands, West Guadalcanal and on Malaita it is 
called either 'ogu' or 'odu' and at the eastern end of San Cristobal it is known as 
'parenga'. Lever (1945) records that it is called 'orku' on Ulawa Island. Collection 
of the 'rising' appears to be a dying custom in the Solomon Islands since many of the 
natives interviewed stated that they had not done so for many years and certainly 
only the older men could relate the details of the event. From their description of 
the rising, it would appear that the spawning of this species appears to be essentially 
similar to that described for other areas. 

The author could not be present at a rising but fortunately a valuable collection of 
the spawning worms was obtained by Wilson Ifunaoa (a schoolboy attached to the 
Expedition) from Fanalei Island (also called Falelei or Halelei Island), off the south- 
east coast of Maramasike Island, Malaita. This collection was made at the time of 
the rising on 2~3.xi.66 and four species are represented - Phyllodoce (Anaitides) 
madeirensis, Perinereis cultrifera, Eunice coccinea and Lysidice collaris, in addition to 
Eunice (Palola) siciliensis. The details of this material, together with the spawning 
times provided by the collector, are given in Table 6. There can be little doubt 
that all five species were spawning at the same time because all of the specimens 
either contain apparently mature gametes or are in the 'spent' condition. In a 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 



119 



similar collection from the New Hebrides, Gravier (1924) records that Lumbrineris 
sphaerocephala and Perinereis masalacensis were present. 

TABLE 6 

Details of the material in the sample of the 'palolo' or 'ogu' rising at Fanalei, Malaita, collected 

on 2-3.xi.66 by Wilson Ifunaoa. 



Species 



Phyllodoce 
(Anaitides) 
madeirensis 

Perinereis 
cultrifera 

Eunice 
coccinea 

Eunice 

(Palola) 

siciliensis 

Lysidice 
collaris 



Material 



3 specimens 



2 heteronereids 



2 posterior 
fragments 

4 posterior 
fragments 



i specimen 



Time of 
spawning (hours) 



1830-1900 



2OOO-22OO 



0000-0600 



2OOO-22OO 



Condition 



Spent 



Ripe oocytes 
200 (A in diameter 

Ripe oocytes 
360 (i in diameter 

Ripe oocytes 
200 (A in diameter 



Spent 



Local name 



Adio 



Falisu-ogu 



Raka-raka 



Falisu-ogu 



Also included in the collection of the 'palolo' or 'ogu' is a large specimen of Notopygos 
gregoryi, a species hitherto recorded only from Midway Island. It is thought that 
this specimen was feeding on the spawning worms as they emerged from the coral. 



SYSTEMATIC ACCOUNT 

The great majority of the records from the littoral zone were made between mid- 
tide level (MTL) and low water mark (LWM). Sublittoral or shallow-water samples 
were taken to a maximum depth of 35 m in Marovo Lagoon. For each species the 
number of specimens taken at each of the localities or stations (see figs i and 2) is 
given : where more than 20 specimens were found the species is indicated as numerous 
(num.) 

Family APHRODITIDAE 

Subfamily POLYNOINAE 

Gastrolepidia clavigera Schmarda, 1861 

Gastrolepidia clavigera Schmarda, 1861 : 159, pi. 36, fig. 316; Fauvel, 1953 : 51, fig. 22. d-f; 
Day, 1967 : 51, fig. 1.5. a-f. 

HABITAT. A commensal of holothurians (see Gibbs, 1969 for notes). 



izo P. E. GIBBS 

RECORDS. On Stichopus chloronotus Brandt - Graham Pt. -8; Maraunibina 
Is. - 12; Kalota Is. - 2; on Holothuria (Halodeima) atra Jaeger - Graham Pt. - u ; 
Kalota Is. - 5; Auki Hr. - num. ; on Bohadschia argus Jaeger - Maraunibina Is. - 4; 
Kalota Is. - 15; on Bohadschia graffei Semper - Maraunibina Is. - 2 ; Yandina - i ; 
on Thelenota ananas (Jaeger) - Maraunibina Is. - I. 

DISTRIBUTION. Tropical Indo-west-Pacific. 



Harmothoe nigricans Horst, 1915 

Harmothoe nigricans Horst, 1915 : 14; 1917 : 90, pi. 20, figs 3-4. 
HABITAT. Under coral boulders and in beachrock. 
RECORDS. Tetel Is. - i ; Komimbo Bay - 4; Lauvie Is. - i. 
DISTRIBUTION. East Indies. 



Harmothoe sp. 

HABITAT. Crevice in reef platform. 
RECORD. Maramasike Pg. - i. 

NOTES. The specimen has the typical harmothoid prostomium with pronounced 
lateral peaks and ventrally inserted lateral tentacles. The elytra number fifteen 
pairs and each carries large conspicuous tubercles. The latter are pear-shaped with 
wide bases, the largest of them being situated along the posterior margins of the 
elytra. Notosetae are numerous and coarsely serrated and neurosetae bidentate 
with a fine secondary tooth almost as long as the main one. 

The specimen resembles H, impar (Johnston), known from Japan (Imajima and 
Hartman, 1964) but differs in that the elytral margins are strongly ciliated. 



Hololepidella nigropunctata (Horst, 1915) 

Polynoe nigro-punctata Horst, 1915 : 20; 1917 : 104, pi. 21, figs 15-17. 

Hololepidella nigropunctata; Day, 1957 : 65, fig. i. a-f; Devaney, 1967 : 287, figs 1-5; Pettibone, 

igdga : 50, fig. 2. a-g. 

Polyeunoa nigropunctata: Day, 1967 : 54, fig. 1.5. r-u. 
Hololepidella minuta: Gibbs, 1969 : 449, fig. 131. 

HABITAT. A commensal of ophiuroids living under coral boulders and in crevices 
on the reef platform (see Gibbs, 1969). 

RECORDS. Graham Pt. - on Ophiocoma brevipes Peters - 8 ; on Ophiocoma insularia 
forma dentata Liitken - i ; on Ophiarthrum elegans Peters - 7. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 121 

NOTES. In the preliminary report on the commensal species (Gibbs, 1969) these 
specimens were referred to Hololepidella minuta (Potts, 1910) because they correspond, 
particularly in terms of their setae, with the type specimen of Polynoe minuta in the 
British Museum (Natural History). The latter specimen is incomplete however, 
having only 24 segments, and thus cannot be referred to the genus Hololepidella 
Willey with certainty because the generically characteristic, segmental arrangement 
of the elytra (Devaney, 1967; Pettibone, 1969^) cannot be checked. Unfortunately 
the type specimens of Polynoe minuta var. oculata Potts (1915) must be considered 
lost since they are not present in the British Museum nor in the Zoological Museum 
of the University of Cambridge (C. B. Goodhart, pers. comm.). Thus it seems 
advisable to refer the Solomon Islands specimens to H. nigropunctata (Horst) and to 
regard P. minuta as indeterminable. Possibly further specimens from crinoids at 
the type localities of P. minuta (South Male) and P. minuta oculata (Torres Straits) 
may clarify the problem. 

DISTRIBUTION. Tropical Indo-west-Pacific to Hawaii. 



Hololepidella ophiuricola n. sp. 

(Fig. 3. A-H) 

Hololepidella commensalis: Gibbs, 1969 : 451, fig. 132. 

DESCRIPTION. The largest specimen (holotype) measures 15 mm in length, is 
2-5 mm wide and has 53 segments bearing 25 pairs of elytra. Smaller specimens 
have 41 to 50 segments bearing 19 to 24 pairs of elytra. 

The prostomium has marked antero-lateral peaks and two pairs of eyes (fig. 3. A). 
The median antenna and tentacular cirri are long and the lateral antennae are short 
and ventral in origin. The palps are stout and between 1-5 to 2-0 times the length 
of the prostomium. Both dorsal and ventral cirri are relatively long, the former 
being about equal to the width of the body, the latter about equal to the length of the 
neuropodial lobe. 

The elytra are oval in shape and completely cover the body. They have smooth 
margins and are transparent except for small, irregular patches of black pigment 
contained in polygonal-shaped cells (fig. 3. B, c). As is characteristic of the genus, 
the elytra are carried on segments 2, 4, 5, 7, 9, . . . 21, 23, 26, 29, 31, 34, 36 and on 
alternate segments to the posterior end. The number of elytra depends on the 
segment number and in four of the five specimens there are 24 or 25 pairs on 49 to 
53 segments. 

The notopodial lobe is rather small (fig. 3. D) and carries 16 to 20 notosetae, each 
of which is minutely serrated along one edge (fig. 3. E). The neuropodium is bilobed 
with a triangular pre-setal lip and a rounded post-setal lip. There are 3 or 4 supra- 
acicular and 6 to 8 sub-acicular neurosetae. Superior neurosetae have about 
15 rows of spinules and a smooth, curved tip (fig. 3. F) : inferior neurosetae are shorter 
and more slender with minute serrations (fig. 3. G, H). 



122 P. E. GIBBS 

All specimens have a darkly pigmented dorsal surface the pattern of which may 
vary according to the species of the host ophiuroid (see Gibbs, 1969). The cirri and 
antennae are dark, as are the palps which also show a white longitudinal line on their 
dorsal surface. Apart from a dark circular spot at the base of each parapodium, the 
ventral surface is unpigmented. 

H. ophiuricola may be distinguished from the type species H. commensalis Willey 
and from H. nigropunctata by its notosetae, the ornamentation of which differs in all 
three species. Also H. ophiuricola differs from the former species in having a greater 
number of notosetae (16 to 20 compared to 8) and from H. nigropunctata in having 
unidentate, not bidentate neurosetae. It should be noted that in a recent review of 
the genus Hololepidella Pettibone (19690) considers the records of H. commensalis 
given by Augener (1922) and Fauvel (1932) from the Indian Ocean are doubtful in 
view of the fact that both sets of specimens differ from the type description in having 
bidentate neurosetae. 



0.5mm 




FIG. 3. Hololepidella ophiuricola n.sp. (A) Dorsal view of anterior region showing 
pigmentation. (B) Elytron from middle body, (c) Patch of pigment cells on elytron. 
(D) Elytrigerous parapodium from middle body. (E) Notoseta. (F-H) Superior and 
inferior neurosetae. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 123 

HABITAT. A commensal of ophiuroids living under coral boulders and in crevices 
on the reef platform. 

RECORDS. Graham Pt. - on Macrophiothrix koehleri Clark - 4 (including holo- 
type) ; Kalota Is. - on Ophiarthrum pictum Miiller and Troschel - i. 
British Museum (Natural History) Registration No. Holotype 1970-20 

Paratypes 1970-21-24 



Iphione muricata (Savigny, 1818) 

Iphione muricata: Fauvel, 1953 : 3 2 > fig- X 3- a ~ e ; Day, 1967 : 43, fig. 1.3. a-f. 
HABITAT. Under coral boulders in muddy silt and sand. 

RECORDS. Tetel Is. - 3; Komimbo Bay - 3; Maraunibina Is. - 2; Graham 
Pt. - 24. 

DISTRIBUTION. Tropical Indo-west-Pacific. 



Lepidasthenia elegans (Grube, 1840) 

Lepidasthenia elegans : Potts, 1910 : 342, pi. 19, fig. 16, pi. 20, fig. 32; Fauvel, 1923 : 88, fig. 33. 
a-g; Day, 1967 : 90, fig. 1.16. i-m. 

HABITAT. In silt and cohabiting a tube of Mesochaetopterus Sagittarius at LWM. 

RECORDS. Tetel Is. - 2. 

DISTRIBUTION. Mediterranean; Indo-west-Pacific. 



Lepidasthenia guadalcanalis n. sp. 

(Fig. 4. A-F) 

Lepidasthenia mossambica: Gibbs, 1969 : 453, fig. 134. 

DESCRIPTION. The holotype is the largest of the six specimens and measures 85 mm 
for about 125 segments. Preserved in alcohol the colour is pale brown with darker 
bars across the anterior segments. The fourth pair of elytra is coloured reddish- 
brown (fig. 4. A) but the other elytra are colourless. 

The prostomium is roughly hexagonal in shape and has two pairs of eyes situated 
on the antero-lateral and posterior margins (fig. 4. B). The median and lateral 
antennae are terminal in origin and are about one half the length of the stout palps. 
The tentacular cirri are slightly longer than the antennae. The posterior margin 
of the prostomium is hidden by a conspicuous occipital flap which is heavily papil- 



I2 4 



P. E. GIBBS 



lated. Similar papillae are to be found along the anterior and posterior edges of the 
anterior segments on their dorsal side and a few are present on the dorsal and ventral 
surfaces of the neuropodial lobes. 

The elytra, which are arranged segmentally in the typical polynoid sequence, are 
thin, circular in shape and do not cover the mid-dorsal region of the body. They 
have smooth margins and lack papillae. Only the fourth pair, on segment 7, is 
pigmented and the others are almost transparent. 

The dorsal cirri are about equal in length to the neuropodial lobes. The noto- 
podial lobes are small and supported by an aciculum but the neuropodial lobes are 
well-developed with rounded, subequal, pre-setal and post-setal lips and short 
ventral cirri (fig. 4. c). 

Notosetae appear to be entirely lacking. In the anterior segments the neurosetae 
are slender with a small subterminal tooth (fig. 4. D, E) and number about 30 per 
neuropodium. However in the middle body segments, the neurosetae are much 
stouter, fewer in number (14 or 15 per neuropodium) and lack subterminal teeth 
(fig. 4. F). The number of rows of spinules possessed by these setae is similar 




0.5mm 

FIG. 4. Lepidasthenia guadalcanalis n.sp. (A) Dorsal view of anterior region showing 
pigmentation of fourth pair of elytra. (B) Head region, (c) Parapodium from middle 
body. (D-E) Superior neuroseta from anterior segment. (F) Superior neuroseta from 
middle body segment. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 125 

throughout the body, the superior ones having about 7, the inferior ones 4 or 5. 
Neither slender superior neurosetae in the anterior segments nor giant neurosetae 
in the posterior region could be found. 

In possessing a distinctly papillated occipital flap, L. guadalcanalis resembles three 
other species, namely, L. michaelseni Augener, L. terrareginae Monro and L. mossambica 
Day. It differs from the former two species in lacking slender superior neurosetae 
and in having unidentate neurosetae in the middle segments. L. guadalcanalis is 
close to L. mossambica, to which these specimens were provisionally assigned (Gibbs, 
1969), but a comparison of the type specimen of L. mossambica (in the British 
Museum, Natural History) has revealed important differences in the structure of the 
setae. In the middle segments of L. guadalcanalis the neurosetae are much stouter, 
fewer in number and have fewer rows of spinules (7 compared to 10 or 12) than in 
L. mossambica and the presence of small subterminal teeth on the neurosetae of the 
anterior segments of L. guadalcanalis also separates the two species. Furthermore 
the distinctive coloration of the fourth pair of elytra appears to be a characteristic 
feature of L. guadalcanalis. 

HABITAT. A commensal of a large enteropneust (Balanoglossus carnosus?} living 
in silty sand (Thalassia flat). 

RECORDS. Graham Pt. - 4; Fintry Pt. - 2 (including holotype). 
British Museum (Natural History) Registration No. Holotype 1970-25 

Paratypes 1970-26-28 



Lepidasthenia maculata Potts, 1910 

Lepidasthenia maculata Potts, 1910 : 344, pi. 20, fig. 33, pi. 21, fig. 51; Fauvel, 1953 : 58, fig. 27. 
h-k; Day, 1967 : 92, fig. 1.16. s-v. 

HABITAT. In coarse sand, silt and within a tube of Phyllochaetopterus herdmani. 

RECORDS. Haroro - i ; Tetel Is. - 2. 

DISTRIBUTION. North Atlantic; Indian Ocean; Indo-China. 



Lepidasthenia microlepis Potts, 1910 

Lepidasthenia microlepis Potts, 1910 : 343, pi. 19, fig. 17, pi. 21, fig. 52; Fauvel, 1953 : 57, fig. 26. 
e-f; Day, 1967 : 90, 1.16. e-h. 

HABITAT. A commensal of sipunculids. Hosts include Siphonosoma vastus 
(Selenka & Biilow) living in silty sand (Thalassia flat) and Paraspidosiphon cumingi 
(Baird) boring in a Porites boulder. 

RECORDS. Graham Pt. - 4. 
DISTRIBUTION. Tropical Indo-west-Pacific. 



126 P. E. GIBBS 

Lepidasthenia stylolepis Willey, 1907 

Lepidasthenia stylolepis Willey, in Lloyd, 1907 : 260, figs 1-4; Gibbs, 1969 : 453, fig. 134. 

HABITAT. A commensal of the sipunculid Siphonosoma vastus, living in silty sand 
(3 out of 7 specimens were discovered in association; hosts were not found for the 
other 4 specimens). 

RECORDS. Graham Pt. - 7. 

NOTES. The similarity of Perolepis regularis Ehlers and Lepidasthenia sibogae 
Horst to L. stylolepis has been referred to in an earlier paper (Gibbs, 1969). 

DISTRIBUTION. Persian Gulf; PEast Africa; PEast Indies. 



Lepidonotus (Thormora) jukesi (Baird, 1865) 

Lepidonotus (Thormora) jukesi: Fauvel, 1953 : 37, fig. 13. o-r; Day, 1967 : 80, fig. 1.13. g-m. 

HABITAT. Under coral boulders, in crevices within reef platform, amongst 
Phyllochaetopterus socialis tubes and algal cover (Amphiroa) from MTL to LWM. 

RECORDS. Tetel Is. - i ; Matiu Is. - 13 ; Batuona Is. - 17 ; New Manra - 5 ; 
Komimbo Bay - i ; Mamara Pt. - I ; Lingatu - 2 ; Maramasike Pg. - 3. 

DISTRIBUTION. Indo-west-Pacific. 



Paradyte crinoidicola (Potts, 1910) 

Polynoe crinoidicola Potts, 1910 : 337, pi. 18, fig. 10, pi. 20, fig. 30, pi. 21, figs 39-41. 
Scalisetosus longicirrus : Fauvel, 1953 : 50, fig. 22. a-c; Day, 1967 : 58, fig. 1.7. a-f. 
Paradyte crinoidicola: Pettibone, 1969 : 13, fig. 7. a-g. 

HABITAT. A commensal of crinoids; on Himerometra robustipinna (P. H. 
Carpenter). 

RECORDS. Maraunibina Is. - 5 specimens from two hosts. 

NOTES. Following Fauvel (1953) and Day (1967) P. crinoidicola was previously 
referred (Gibbs, 1969) to S. longicirrus (Schmarda) which Pettibone (1969) considers 
indeterminable. 

DISTRIBUTION. Indo-west-Pacific. 



Paralepidonotus ampulliferus (Grube, 1878) 

Polynoe ampullifera Grube, 1878 : 35, pi. 3, fig. 5. 

Harmothoe ampullifera: Fauvel, 1953 : 43, fig. i8.d; Pillai, 1965 : 117, fig. 3. d-g, fig. 4. a-b. 

Paralepidonotus ampulliferus: Horst, 1917 : 76; Day, 1967 : 47, fig. 1.4. a-f. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 127 

HABITAT. Undersurface of coral boulder on reef platform. 

RECORD. Kalota Is. - i. 

DISTRIBUTION. Tropical Indo-west-Pacific. 



Paralepidonotus indicus (Potts, 1910) 

Lagisca indica Potts, 1910 : 338, pi. 19, fig. 13, pi. 21, figs 46-47. 
Paralepidonotus indicus: Day, 1967 : 48, fig. 1.4. g-k. 

HABITAT. Silty sand with coral debris at LWM. 
RECORD. Komimbo Bay- i. 
DISTRIBUTION. Mogambique; Maldive Is. 



Subfamily POLYODONTIDAE 
Polyodontes maxillosus (Ranzani, 1817) 

Polyodontes maxillosus: Fauvel, 1923 : 97, fig. 37. a-n; 1953 : 71, fig. 32. a-n. 

HABITAT. Silty sand with shell and coral fragments at LWM; coarse sand at 
1 8 m depth. 

RECORDS. Graham Pt. - i ; Fintry Pt. - i ; ML 204 - i. 

NOTES. Fauvel records that this species may grow to i m in length. The largest 
of the three specimens from the Solomon Islands measures only 12 cm: its felt-like 
tube had a diameter i-o to 1-5 cm and extended to a depth of about 30 cm into the 
deposit at Graham Pt. 

DISTRIBUTION. North Atlantic ; Mediterranean ; Indian Ocean ; Indo-China. 



Polyodontes melanonotus (Grube, 1876) 

Polyodontes melanonotus: Fauvel, 1953 : 72, fig. 33. c-g; Day, 1967 : 96, fig. 1.17. g-n. 

HABITAT. Mud at n m; coarse sand at 18 m. 
RECORDS. ML 194 - i ; ML 204 - i. 

NOTES. Specimen from ML 194 is 3-5 cm long: it was removed from a tube 
about 15 cm in length, i-o cm in diameter, composed of consolidated mud. 

DISTRIBUTION. West Africa; Indo-west-Pacific. 



128 P. E. GIBBS 

Subfamily SIGALIONINAE 
Psammolyce zeylanica Willey, 1905 

Psammolyce zeylanica Willey, 1905 : 255, pis 1-2, figs 33-43; Fauvel, 1953 : 68, fig 31.!. 
HABITAT. Silty sand (Thalassia flat). 
RECORDS. Graham Pt. - 2. 
DISTRIBUTION. Ceylon. 

Sigalion bandaensis Horst, 1917 

Sigalion bandaensis Horst, 1917 : no, pi. 22, figs 4-5. 

HABITAT. Under boulder on silty sand with coral debris at LWM. 

RECORD. Graham Pt. - i. 

DISTRIBUTION. East Indies; East Australia. 



Sthenelais heterochela Horst, 1917 

Sthenelais heterochela Horst, 1917 : 113, pi. 23, figs 3-6. 

HABITAT. Mud and silty sand, 2-11 m depth. 
RECORDS. ML 191 - i ; ML 194 - i ; ML 195 - i. 
DISTRIBUTION. East Indies. 



Sthenelais zeylanica Willey, 1905 

Sthenelais zeylanica Willey, 1905 : 258, pi. 2, fig. 48; Fauvel, 1953 : 62, fig. 29. a. 
HABITAT. Silty sand, MTL to LWM ; sand, 5-16 m depth. 

RECORDS. Tetel Is. -i; Komimbo Bay -4; Graham Pt. -2; Fintry Pt. -3; 
ML 134 - 2 ; ML 190 - i. 

NOTE. In addition to the two slender stylodes arising from the base of the ventral 
cirrus, these specimens have a third appendage with a bulb-shaped ending. 

DISTRIBUTION. India; Palau Islands. 



Sthenelanella ehlersi (Horst, 1916) 

Euleanira ehlersi Horst, 1916 : 12; 1917 : 122, pi. 27, figs 1-5; Day, 1967 : 101. 
Sthenelanella ehlersi: Pettibone, 19696 : 434, figs 4-5. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 129 

HABITAT. Mud and silty sand, 18-24 m depth. 
RECORDS. ML 188 - 3; ML 230- i. 
DISTRIBUTION. Natal; East Indies. 



Sthenolepis japonica (Mclntosh, 1885) 

Leanira japonica Mclntosh, 1885 : 154, pi. 22, fig. 3, pi. I4A, figs 1-2; Fauvel, 1953 : 69, fig. 

33-a-b. 
Sthenolepis japonica: Imajima & Hartman, 1964 : 43; Day, 1967 : 112. 

HABITAT. A characteristic species of fine deposits in the sublittoral zone, thick 
mud to silty sand, 2-33 m depth. 

RECORDS. ML 56 - 2 ; ML 69 - i ; ML 100 - 5 ; ML 155 - 2 ; ML 156 - 2 ; ML 157 - 
i; ML 188-2; ML 190 -2; ML 191-1; ML 195 -3; ML 196 -3; ML 218 -4; 
ML 228 - i. 

DISTRIBUTION. Indo-west-Pacific to Japan. 



Thalenessa digitata Mclntosh, 1885 

Thalenessa digitata Mclntosh, 1885 : 140, pi. 22, fig. 2, pi. 23, figs 5-7, pi. 25, figs 4-5, pi. I3A, 
figs 7-10; Willey, 1905 : 260, pi. 2, figs 50-52; Imajima & Hartman, 1964 : 46. 

HABITAT. Coarse sand, 2-4 m depth. 
RECORDS. ML 203 - 2 ; ML 296 - i. 
DISTRIBUTION. Ceylon ; Admiralty Is. ; Japan. 



Family PALMYRIDAE 
Bhawania goodei Webster, 1884 

Bhawania cryptocephala Gravier, 1901 : 263, pi. 10, figs 152-156, text-figs 280-285; Fauvel, 

1953 : 79, ng. 36. e-i. 
Bhawania goodei: Day, 1953 : 407 (synonymy); 1967 : 118, fig. 2.1. a-f. 

HABITAT. Cohabiting burrows of coral-boring sipunculids; hosts include Aspido- 
siphon elegans (Chamisso & Eysenhardt), Cloeosiphon aspergillum (Quatrefages) and 
Phascolosoma albolineatum (Baird). Also found free-living in crevices and amongst 
tubes of Phyllochaetopterus socialis. 

RECORDS. Matiu Is. - 30; New Manra - 5 ; Mamara Pt. - i ; Maramasike Pg. - 2. 

DISTRIBUTION. Circumtropical. 



130 P. E. GIBBS 

Bhawania pottsiana Horst, 1917 

Bhawania cryptocephala: Potts, 1910 : 328. 

Bhawania cryptocephala var. pottsiana Horst, 1917 : 137. 

Bhawania sp.: Gibbs, 1969 : 454. 

HABITAT. Sedimented crevices in Porites boulders; commensal (?) with Eurythoe 
complanata (see Gibbs, 1969). 

RECORDS. Graham Pt. - 2. 

NOTES. The larger of the two specimens measures 45 mm in length. Both 
possess very slender setae with hair-like appendices in the inferior part of the 
neuropodium, which are not present in B. goodei. This additional type of seta was 
first noticed by Potts (1910) in a specimen from Zanzibar which he attributed to 
B. cryptocephala. However Horst (1917), in recording a similar specimen from the 
Celebes Is., gave this form varietal status, naming it B. cryptocephala var. pottsiana. 
Day (1953) has shown B. cryptocephala to be a synonym of B. goodei, but since the 
variety pottsiana is quite distinct from its stem species, its subspecific status is here 
raised to specific rank. 

DISTRIBUTION. Zanzibar; East Indies. 



Paleanotus debilis (Grube, 1855) 
Paleanotus debilis: Day, 1962 : 635 (synonymy); 1967 : 117, fig. 2.1. g-k. 

HABITAT. Amongst Phyllochaetoptems socialis tubes at LWM; on Halichondria 
sp. at 5 m depth. 

RECORDS. Matiu Is. - i ; Batuona Is. - i ; Yandina - 2. 
DISTRIBUTION. North Atlantic; Mediterranean; Indo-Pacific. 



Family AMPHINOMIDAE 
Amphinome nigrobranchiata Horst, 1912 

Amphinome nigrobranchiata Horst, 1912 : 39, pi. 10, figs 17-20. 
HABITAT. Under coral boulders; in shell gravel deposits. 
RECORDS. Tetel Is. - 2 ; Graham Pt. - 2 ; Maraunibina Is. - i. 
DISTRIBUTION. East Indies. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 131 

Chloeia conspicua Horst, 1910 

Chloeia conspicua Horst, 1910 : 173; 1912 : 20, pi. 7, fig. 5, pi. 8, figs 4-5. 

HABITAT. Silty sand at 2 m depth. 
RECORD. ML 196 - i. 
DISTRIBUTION. East Indies. 

Euphrosine foliosa Audouin & Milne-Edwards, 1833 

Euphros ine foliosa : Fauvel, 1923 : 136, fig. 49. a-g; 1953 : 102, fig. 48. a-h. 
HABITAT. Under coral boulder at LWM. 
RECORD. Maraunibina Is. - i. 
DISTRIBUTION. Atlantic Ocean; Mediterranean; Indo-west-Pacific. 

Euphrosine myrtosa Savigny, 1818 

Euphrosine myrtosa : Fauvel, 1923 : 139, fig. 49. k-n; 1953 : 101, fig. 48. k-n; Day, 1967 : 127, 
fig. 3.1.2. 

HABITAT. Amongst Phyllochaetopterus socialis tubes; sedimented crevice in 
Porites boulder. 

RECORDS. Batuona Is. - i ; Graham Pt. - i. 

DISTRIBUTION. West Africa; Mediterranean; Indo-west-Pacific. 



Eurythoe complanata (Pallas, 1766) 
Eurythoe complanata : Fauvel, 1953 : 83, fig. 38. b-m; Day, 1967 : 128, fig. 3.2. a-h. 

HABITAT. A wide variety of situations, including under coral boulders, in crevices 
in reef platform and in beachrock, in Acropora rubble and amongst Phyllochaetopterus 
socialis tubes and Amphiroa. 

RECORDS. Tetel Is. -10; Komimbo Bay -20; Matiu Is. -num. ; Batuona Is. - 
num. ; New Manra - i ; Graham Pt. - 6; Maraunibina Is. - 5 ; Lauvie Is. - i. 

DISTRIBUTION. Circumtropical. 



? Eurythoe sp. 

HABITAT. Under coral boulders in silty sand. 
RECORDS. Maraunibina Is. - 4. 

NOTES. The specimens measure between 10 and 20 mm long and about 2 mm 
wide and have 57 to 67 segments. A sinuous caruncle extends through setiger 2 



132 P. E. GIBBS 

and the branchiae commence between setigers 9 and 13, extending to the posterior 
end. Notosetae are of three types, namely (i) stout spines, (ii) harpoon setae and 
(iii) long, fine, smooth capillaries. Neurosetae are of two types - (i) furcate, with a 
short secondary spur and (ii) long, fine capillaries with faintly serrated blades, some 
with a stout spur. 

The generic identity of these specimens is in question since in the genus Eurythoe 
the branchiae commence on setigers 1-3 (Day, 1967). It is possible that these 
relatively small specimens are juveniles and because of this, the erection of a new 
genus must be postponed until further material is available. 



Notopygos gregoryi Holly, 1939 

Notopygos gregoryi Holly, 1939 : 265, fig. i. a-f. 

HABITAT. Not known. 
RECORD. Fanalei - 1. 

NOTES. The specimen of this elegant species was collected during the 'palolo' 
rising at Fanalei between 2000 and 2200 hours on 2.xi.66. It is 160 mm long and 
25 mm across the body. All details correspond to the description of the holotype. 

DISTRIBUTION. Midway Is. (one specimen). 



Notopygos sibogae Horst, 1911 

Notopygos sibogae Horst, 1911 : 245; 1912 : 27, pi. 9, figs 4-5. 

HABITAT. In Acropora rubble ; within sedimented crevices on the undersurfaces 
of Porites boulders. 

RECORDS. Tetel Is. - i ;, Graham Pt. - 3. 

NOTES. Hartman (1966) comments that the differences in the specific characters 
in Notopygos i.e. number of segments and the position of the anal pore, may reflect 
differences in the growth stages. However in all four specimens in the collection of 
N. sibogae the anal pore is situated on the anterior margin of setiger 23 although the 
lengths vary from 10 to 20 mm. 

DISTRIBUTION. East Indies. 



Pareurythoe pitipanaensis De Silva, 1965 

Pareurythoe pitipanaensis De Silva, 1965 : 540, fig. 3. a-k. 

HABITAT. Under coral boulders, in shell gravel and within the interstices of the 
coral Galaxea. 

RECORDS. Tetel Is. - i ; Graham Pt. - 5 ; Maraunibina Is. - 3. 
DISTRIBUTION. Ceylon. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 133 

Pseudeurythoe paucibranchiata Fauvel, 1932 

Pseudeurythoe paucibranchiata Fauvel, 1932 : 47, pi. i, figs 3-4, text-fig. 8. a-e; 1953 : 86, fig. 
39. a-b, fig. 40. a-e. 

HABITAT. In silty deposits, particularly under coral boulders; in Acropora 
rubble ; in muds and fine sands, 2-26 m depth. 

RECORDS. Tetel Is. - 3 ; Komimbo Bay - 12 ; Graham Pt. - 7 ; Maraunibina Is. - 
4; ML 37-2; ML 41 - i; ML 68- i; ML 96-1; ML no- 15; ML 134- i; ML 156- 
3 ; ML 157 - 6 ; ML 194 - i ; ML 195 - i ; ML 218 - i ; ML 228 - 4. 

NOTES. Most of the specimens show a button-like caruncle set into the first setiger 
but some are so contracted that this feature is not visible. Small specimens about 
5 mm in length have 4 or 5 pairs of branchiae, whilst in medium-sized specimens up 
to 10 mm long, the branchiae continue to setiger 16 or 17. The largest specimens 
come from the littoral zone (Marau Sound) ; these measure 35 mm long and have 
branchiae up to setiger 27, as typical for P. paucibranchiata. 

There is some doubt as to whether all of the specimens in this series should be 
referred to P. paucibranchiata since the position of the prostomium in relation to the 
first segments and the presence of a caruncle, two major specific characters, are 
difficult to determine in strongly contracted specimens. Further, the number of 
branchiae varies greatly with size and cannot be regarded as specific, particularly in 
small specimens. The setae also are not diagnostic, according to Wesenberg-Lund 
(1949). On account of these considerations, it seems probable that a number of 
Pseudeurythoe species are synonyms of P. oculifera (Augener) , the description of which 
was based on a small, probably juvenile, specimen. 

DISTRIBUTION. Indian Ocean ; Indo-China. 



Family PHYLLODOGIDAE 
Eteone japanensis Mclntosh, 1901 

Eteone japanensis Mclntosh, 1901 : 222, pi. i, fig. 2; Imajima & Hartman, 1964 : 60., 

HABITAT. Silty sand. 
RECORD. Tetel Is. - i. 

NOTES. The specimen agrees with those details given in the type description and, 
in addition, possesses a smooth proboscis. However, since the characters of the 
proboscis in this species are unknown the identification must be regarded as a 
provisional one. 

DISTRIBUTION. Japan Sea. 



134 p - E - GIBBS 

Eulalia albopicta Marenzeller, 1879 

Eulalia albopicta Marenzeller, 1879 : 128, pi. 3, fig. 3; Fauvel, 1953 : 123, fig. 60. a-b. 
HABITAT. Cohabiting a Phyllochaetopterus elioti tube from silty sand. 
RECORD. Fintry Pt. - i. 
DISTRIBUTION. Indian Ocean; Indo-China; Japan. 

Eulalia viridis (Linnaeus, 1767) 

Eulalia viridis : Fauvel, 1923 : 160, fig. 57. a-h; 1953 : 122, fig. 6.1. a-h; Imajima & Hartman, 
1964 : 63. 

HABITAT. Crevice in reef platform. 
RECORD. Maramasike Pg. - i. 
DISTRIBUTION. Cosmopolitan. 

Eulalia (Pterocirrus) magalhaensis Kinberg, 1866 

Eulalia (Pterocirrus) magalhaensis : Kinberg 1858-1910 : 55, pi. 33, fig. i; Fauvel, 1953 : 124, 
fig. 62. a-h. 

HABITAT. Amongst Phyllochaetopterus socialis tubes and algal growth (Amphiroa) 
towards LWM. 

RECORDS. Cape Esperance - i ; Matiu Is. - 2; Batuona Is. - i ; New Manra - i. 

NOTE. All five specimens are small-sized (3 to 7 mm) but conform to the species 
description. 

DISTRIBUTION. Indo-Pacific. 

Notophyllum splendens (Schmarda, 1861) 

Macrophyllum splendens Schmarda 1861 : 82, pi. 29, fig. 227. 

Notophyllum splendens: Fauvel, 1953 : 126, fig. 60. c; Day, 1967 : 151, fig. 5.3. k-n. 

HABITAT. Under coral boulder; interstices of the coral Galaxea. 
RECORDS. Graham Pt. - 2. 
DISTRIBUTION. Indo-west-Pacific. 

Phyllodoce fristedti Bergstrom, 1914 

Phyllodoce fristedti : Fauvel, 1953 : 118, fig. 58. a-b; Day, 1967 : 147, fig. 5.2. k-m. 

HABITAT. Crevices in reef platform and beachrock. 
RECORDS. Matiu Is. - i ; Lauvie Is. - i. 
DISTRIBUTION. Indian Ocean. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 135 

Phyllodoce malmgreni Gravier, 1900 

Phyllodoce malmgreni Gravier, 1900 : 207, pi. 10, figs 2931, text-figs 66-69; Fauvel, 1953 : JI 7 
fig. 56. h; Day, 1967 : 147, fig. 5.2. n-p. 

HABITAT. Silty sand at MTL; mud at 2 m depth. 
RECORDS. Tetel Is. - 2 ; ML 68 - i. 
DISTRIBUTION. Indian Ocean; East Australia. 



Phyllodoce pruvoti Fauvel, 1930 
Phyllodoce pruvoti Fauvel 1930 : 512, fig. i. a-f, fig. 2. a-c; 1947 : 28, fig. 23. 

HABITAT. Crevices in reef platform ; amongst Phyllochaetopterus socialis tubes. 
RECORDS. Matiu Is. - i ; Batuona Is. - i ; New Manra - i. 
DISTRIBUTION. Tropical West Pacific. 



Phyllodoce quadraticeps Grube, 1878 

Phyllodoce quadraticeps Grube, 1878 : 98, pi. 6, fig. 2; Fauvel, 1953 : 116, fig. 56. f-j; Day, 1967 : 
145, fig. 5.2. h-j. 

HABITAT. Crevices in reef platform and in beachrock, MTL to LWM. 
RECORDS. Kokomtambu Is. - i ; Matiu Is. - 10; Lauvie Is. - i. 

NOTE. This species is often found crawling over the surface of exposed coral 
during the low-tide period. 

DISTRIBUTION. Tropical Indo-west-Pacific. 



Phyllodoce (Anaitides) madeirensis Langerhans, 1880 

Phyllodoce (Anaitides) madeirensis: Fauvel, 1953 : 120, fig. 59. d-h; Day, 1967 : 145, fig. 5.2. d-g. 

HABITAT. In Acropora rubble, under coral boulders and in beachrock crevices. 
RECORDS. Komimbo Bay - 2 ; Lauvie Is. - 2 ; Graham Pt. - 2 ; Fanalei - 3. 
NOTES. The Fanalei specimens were collected between 1830 and 1900 hours 
during the 'palolo' rising on 2.xi.66 and are 'spent'. Local name is 'adio'. 

DISTRIBUTION. Cosmopolitan. 



136 P. E. GIBBS 

Phyllodoce (Anaitides) parva (Hartmann - Schroder, 1965) 
Anaitides parva Hartmann - Schroder, 1965 : 88, figs 7-9; Hartman, 1966 : 183. 

HABITAT. Medium sand at 5 m. 
RECORD. ML 190-1. 
DISTRIBUTION. Hawaii. 



Phyllodoce (Genetyllis) castanea (Marenzeller, 1879) 

Carobia castanea Marenzeller, 1879 : 127, pi. 3, fig. 2; Izuka, 1912 : 199, pi. 21, fig. 3. 
Phyllodoce (Genetyllis) castanea: Fauvel, 1953 : 115, fig. 56. a-c; Day, 1967 : 149, fig. 53. d-f. 

HABITAT. Sand at n m. 
RECORD. ML 98-1. 
DISTRIBUTION. Indo-west-Pacific. 



Phyllodoce (Genetyllis) gracilis Kinberg, 1866 

Phyllodoce gracilis : Kinberg, 1858-1910 : 55, pi. 22, fig. 3; Monro, 1939 : 173, fig. 3. a-c; Fauvel, 
1953 : 117. fi g- 57- a-g. 

HABITAT. Crevice in reef platform. 
RECORD. Lingatu - 1. 
DISTRIBUTION. Tropical Indo-Pacinc. 



Family PILARGIDAE 
Sigambra hanaokai (Kitamori, 1960) 

Ancistrosyllis hanaokai Kitamori, i96oa : 1086, fig. i. a-h. 
Sigambra hanaokai: Pettibone, 1966 : 181 (synonymy). 

HABITAT. Mud, 2-20 m depth. 

RECORDS. ML 39 - 3 ; ML 40 - i ; ML 41 - 10 ; ML 68 - 5 ; ML 157-1. 

NOTES. Larger specimens - up to 14 mm in length - were found inhabiting thin 

membraneous tubes covered in mud particles. 

DISTRIBUTION. Japan. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 137 

Synelmis albini (Langerhans, 1881) 

Ancistrosyllis rigida Fauvel, 1919 : 337, fig. i. a-e; 1953 : no, fig. 53. a-e; Day, 1967 : 215. 
Synelmis albini: Pettibone, 1966 : 191, figs 19-21 (synonymy). 

HABITAT. Crevices in reef platform; amongst Phyllochaetopterus socialis tubes; 
in shell gravel and coarse sand. 

RECORDS. Komimbo Bay - 2 ; Matiu Is. - i ; Batuona Is. - 4; Mamara Pt. - I ; 
Graham Pt. - i ; Maramasike Pg. - i. 

DISTRIBUTION. Circumtropical. 



Family HESIONIDAE 
Gyptis capensis (Day, 1963) 

Oxydromus capensis Day, 1963 : 397, fig. 4. e-j. 
Gyptis capensis: Day, 1967 : 231, fig. 11.2. l-o. 

HABITAT. Under coral boulders on silty sand, MTL to LWM. 
RECORDS. Maraunibina Is. - 3. 

NOTES. A complete specimen measures 15 mm in length for about 70 segments. 
Notosetae are present from setiger 5. 

DISTRIBUTION. South Africa. 



Gyptis maraunibinae n. sp. 

(Fig. 5. A-F) 

DISTRIBUTION. The holotype is 45 mm long for 150 segments and the body is 
about 3 mm wide. 

The prostomium is rectangular, its width being about 1-5 times the length (fig. 5. A). 
Three antennae arise from the anterior margin of the prostomium; the laterals 
are about twice the length of the median. The two biarticulate palps, each com- 
posed of a short palpostyle and longer palpophore, are stouter and slightly longer 
than the lateral antennae. There are two pairs of eyes and those of the anterior 
pair are reniform in shape and are larger than those of the posterior pair. The 
proboscis carries ten papillae, each of which is broad and flap-like (fig. 5. B). Jaws 
are lacking, but there is a horny rim around the ventral and ventro-lateral margins 
of the proboscis. 

The tentacular cirri number eight pairs and each is faintly but closely annulated. 
The dorsal pair of the second segment is the longest, reaching back to about 
setiger 15. 



138 P. E. GIBBS 

The first 4 or 5 setigers are uniramous, becoming biramous when a notopodial 
papilla appears on the antero-ventral side of the cirrophore from about setiger 6. 
The notopodia are small throughout the length of the body but the neuropodia are 
well developed, each with a pointed pre-setal lip and a rounded post-setal lip (fig. 5. c). 
Ventral cirri are about equal to the neuropodial lobe in length. In the middle 
of the body, notosetae include 2 or 3 rather stout, smooth spines (fig. 5. E) and 4 or 
5 forked setae (fig. 5. D) ; neurosetae are falcigerous, each with a minutely serrated 
blade of varying length and with a bidentate tip consisting of a strong terminal 
tooth and a slender secondary one (fig. 5. F). 

The large, lobular, papillae on the proboscis are sufficient to distinguish G. 
maraunibinae from other Gyptis species. In other details the species resembles G. 
capensis (Day). 

HABITAT. Silty sand with shell gravel below coral boulder on reef platform. 

RECORD. Graham Pt. - i. 

British Museum (Natural History) Registration No. Holotype 1970-29 





1.0mm 



0.5mm 




=1 
o 

10 



0.5mm 

FIG. 5. Gyptis maraunibinae n.sp. (A) Dorsal view of prostomium. (B) Anterior view of 
proboscis, (c) Parapodium from middle body. (D-E) Notopodial forked seta and spine. 
(F) Neuropodial falciger. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 139 

Hesione splendida Savigny, 1818 

H esione pantherina : Fauvel, 1953 : IO 4' fig- 49- a "g- 
Hesione splendida: Day, 1967 : 228, fig. 11.2. a-c. 

HABITAT. In Acropora rubble and shell gravel; in crevices and under coral 
boulders. 

RECORDS. Honiara - i ; Komimbo Bay - i ; Graham Pt. - 14. 
DISTRIBUTION. North Atlantic ; Mediterranean ; Indo-west-Pacific. 



Leocrates chinensis Kinberg, 1866 

Leocrates claparedii: Fauvel, 1953 : 106, fig. 50. c-g; Day, 1967 : 230, fig. 11.2. g-k. 
Leocrates chinensis: Imajima & Hartman, 1964 : 82 (synonymy). 

HABITAT. In shell gravel and Acropora rubble; under coral boulders. 

RECORDS. Kokomtambu Is. - i ; Tetel Is. - 4; Komimbo Bay - i ; Fintry Pt. - 
i ; Maramasike Pg. - i. 

DISTRIBUTION. West Africa; Mediterranean; tropical Indo-west-Pacific. 



Family SYLLIDAE 

The species listed below for this family were identified by Dr. Minoru Imajima. 
A detailed account of this material will be published by Dr. Imajima at a later date. 



Subfamily AUTOLYTINAE 
Autolytus sp. 

HABITAT. On sponge Halichondria sp. at 5 m depth. 
RECORD. Yandina- i (damaged). 

Subfamily EXOGONINAE 
Brania clavata (Claparede, 1863) 

Grubea clavata: Fauvel, 1923 : 296, fig. 114 a-e. 
Brania clavata: Imajima, 1966 : 393, fig. i a-g. 

HABITAT. Coarse sand at LWM. 

RECORD. Komimbo Bay - i. 

DISTRIBUTION. North Atlantic; Mediterranean; Japan. 



i 4 o P. E. GIBBS 

Exogone gemmifera Pagenstecher, 1862 

Exogone gemmifera : Fauvel, 1923 : 305, fig. 117. a-d; Imajima, 1966 : 397, fig. 2. a-h; Day, 1967: 
274, fig. 12. 10. p-u. 

HABITAT. Amongst Phyllochaetopterus socialis tubes at LWM. 
RECORD. Matiu Is. - i. 

DISTRIBUTION. North Atlantic; South Africa; Mediterranean; Japan; northern 
North Pacific. 



Exogone uniformis Hartman, 1961 

Exogone uniformis Hartman, 1961 : 73, pi. 6, fig. i, pi. 7, figs 1-4; Imajima, 1966 : 400, fig. 4. 
a-j. 

HABITAT. Coarse sand and silty fine coral sand at LWM. 
RECORDS. Komimbo Bay - i ; Matiu Is. - i. 
DISTRIBUTION. Southern California; Japan. 



Exogone verugera (Claparede, 1868) 

Exogone verugera: Fauvel, 1923 : 307, fig. 117. m-r; Imajima, 1966 : 399, fig. 3. a-h; Day, 1967 : 
272, fig. 12. 10. g-1. 

HABITAT. On Eunice tubifex tube at LWM; on sponge Halichondria sp. at 5 m 
depth. 

RECORDS. Tetel Is. - i ; Yandina - i. 

DISTRIBUTION. North Atlantic; Mediterranean; South Africa; S.W. and E. 
Australia; North Pacific. 



Sphaerosyllis hirsuta Ehlers, 1897 

Sphaerosyllis hirsuta Ehlers, 1897 : 48, pi. 3, figs 58-60; Imajima & Hartman, 1964 : 117, pi. 27, 
figs f-1; Imajima, 1966 : 404. 

HABITAT. Coarse sand at LWM ; on Eunice tubifex tube at LWM. 
RECORDS. Komimbo Bay - i ; Tetel Is. - i. 

DISTRIBUTION. Northern North Pacific; Japan; Australia; New Zealand; 
southern South America. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 141 

Subfamily SYLLINAE 
Haplosyllis spongicola (Grube, 1855) 

Syllis (Haplosyllis) spongicola: Fauvel, 1923 : 257, fig. 95. a-d; 1953 : 147, fig. 75. a-d; Day, 

1967 : 240, fig. 12. i. e-i. 
Haplosyllis spongicola: Imajima, i9&6a : 220, fig. 38. a-h. 

HABITAT. Abundant in sponges - Neofolitispa dianchora and Halichondria sp. - 
from 5 m depth. 

RECORDS. Yandina - num. 

DISTRIBUTION. Cosmopolitan in tropical and temperate waters. 



Langerhansia cornuta (Rathke, 1843) 

Syllis (Ehlersia) cornuta: Fauvel, 1923 : 267, fig. 100 g-i; 1953 : 153, fig. 79. g-i. 
Syllis (Langerhansia) cornuta: Day, 1967 : 244, fig. 12.2. s-u. 
Langerhansia cornuta: Imajima, i966b : 256, fig. 51. a-o. 

HABITAT. Coarse sand, silt ; on sponge Halichondria sp. at 5 m depth. 
RECORDS. Tetel Is. - 3; Komimbo Bay - 19; Yandina - i. 
DISTRIBUTION. Atlantic; Indo-Pacific. 



Langerhansia Irosea (Langerhans, 1879) 
Langerhansia rosea: Imajima, I966b : 259, fig. 52. a-m. 

HABITAT. On serpulid tubes at MTL; coarse sand at LWM. 
RECORDS. Tetel Is. - i ; Komimbo Bay - 2. 



Syllis longissima Gravier, 1900 
Syllis longissima Gravier, 1900 : 154, pi. 9, fig. 7, text-figs. 17-23; Day, 1967 : 243, fig. 12.2. f-i 

HABITAT. Crevices in reef platform ; amongst Phyllochaetopterus socialis tubes at 
LWM. 

RECORDS. Matiu Is. - 8 ; New Manra - i ; Lingatu - 3. 

NOTE. This appears to be the first record of S. longissima in the west Pacific 
region. 

DISTRIBUTION. Red Sea; Persian Gulf; west coast of South America. 



142 P. E. GIBBS 

Syllis sp. cf. gracilis Grube, 1840 

Syllis gracilis: Fauvel, 1923 : 259, fig. 96. f-i; 1953 : 147, fig. 73. f-i; Imajima, ig66a : 248, 
fig. 49. a-k; Day, 1967 : 241, fig. 12.1. m-p. 

HABITAT. Amongst Phyllochaetopterus socialis tubes at LWM; on sponge 
Halichondria sp. at 5 m depth. 

RECORDS. Batuona Is. - 3 ; Yandina - i. 



Trypanosyllis coeliaca Claparede, 1868 

Trypanosyllis coeliaca: Fauvel, 1923 : 270, fig. 101. f-h. 

HABITAT. Amongst Phyllochaetopterus socialis tubes at LWM. 
RECORD. Batuona Is. - i. 

NOTE. Previous records of T. coeliaca are confined to western Europe and the 
Mediterranean Sea. 

DISTRIBUTION. North Atlantic; Mediterranean. 



Trypanosyllis ( Trypanedenta) sp. 

HABITAT. Crevices in beachrock at LWM. 
RECORDS. Lauvie Is. - 3. 



Typosyllis alternata (Moore, 1908) 

Typosyllis alternata: Imajima, I966b : 273, fig. 58. a-1. 

HABITAT. Chiefly amongst Phyllochaetopterus socialis tubes ; under boulders and 
in crevices in reef platform ; on sponge Neofolitispa dianchora at 5 m depth. 

RECORDS. Tetel Is. -2; Matiu Is. -5; Cape Esperance-3; Batuona Is. -6; 
Yandina - 3. 

DISTRIBUTION. Japan; East Indies; west coast of North America. 



Typosyllis armillaris (Miiller, 1776) 

Syllis (Typosyllis) armillaris: Fauvel, 1923 : 264, fig. 99. a-f; Day, 1967 : 249, fig. 12.4. a-d. 
HABITAT. Amongst Phyllochaetopterus socialis tubes at LWM. 
RECORDS. Batuona Is. - 2. 
DISTRIBUTION. Cosmopolitan. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 143 

Typosyllis brachycola (Ehlers, 1897) 

Syllis brachycola Ehlers, 1897 : 38, pi. 2, figs 46-47. 
Syllis (Typosyllis) brachychola: Augener, 1924 : 362. 

HABITAT. Below boulders in silty sand ; amongst Phyllochaetopterus socialis tubes ; 
interstices of coral Galaxea. 

RECORDS. Tetel Is. - i ; Naro Bay - 2; Batuona Is. - 3; Graham Pt. - 2. 

NOTE. T. brachycola is widely distributed in the southern temperate and sub- 
antarctic regions but this appears to be the first record from tropical waters. 

DISTRIBUTION. Southern South America; New Zealand. 



Typosyllis exilis (Gravier, 1900) 

Syllis (Typosyllis) exilis Gravier, 1900 : 160, pi. 9, fig. 9, text-figs 28-30; Day, 1967 : 250, fig. 
12.4. h-j. 

HABITAT. Crevice in reef platform at MTL. 

RECORD. Mamara Pt. - i. 

DISTRIBUTION. Indo-west-Pacific to Gambier Islands. 



Typosyllis lucida (Chamberlin, 1919) 

Pionosyllis lucida Chamberlin, 1919 : 8. 
Typosyllis lucida: Hartman, 1968 : 489. 

HABITAT. Crevices in reef platform. 
RECORDS. Kokomtambu Is. - i ; Matiu Is. - 2. 

NOTE. This species has not been recorded since it was first described by Chamber- 
lin from Laguna Beach, California. 

DISTRIBUTION. Southern California. 



Typosyllis prolifera (Krohn, 1852) 

Syllis (Typosyllis) prolifera: Fauvel, 1923 : 261, fig. 97. a-g; 1953 : 149, fig. 74. a-g; Day, 1967 : 

248, fig. 12.3. g-i. 
Typosyllis prolifera: Imajima, I9&6b : 292, fig. 65. a-n. 

HABITAT. Amongst Phyllochaetopterus socialis tubes at LWM ; on sponge Hali- 
chondria sp. at 5 m depth. 

RECORDS. Batuona Is. - i ; Yandina - 2. 

DISTRIBUTION. North Atlantic; Mediterranean; South Africa; Indo-west-Pacific. 



144 P. E. GIBBS 

Typosyllis sp. 

HABITAT. On sponge Halichondria sp. at 5 m depth. 
RECORD. Yandina - i. 

Family SPHAERODORIDAE 
Sphaerodoridium claparedii (Greeff, 1866) 

Sphaerodorum claparedii: Fauvel, 1923 : 379, fig. 149. d-e. 
Sphaerodoridium claparedii: Liitzen, 1961 : 415. 

HABITAT. On sponge Halichondria at 5 m depth. 
RECORD. Yandina - i. 

NOTES. This specimen was overlooked during the primary sorting of a collection 
of polychaetes from sponges and was included in a large sample of syllids sent to 
Dr. M. Imajima who kindly identified it. This record considerably extends the 
known distribution of 5. claparedii which hitherto was recorded only from western 
Europe. 

DISTRIBUTION. European waters. 



Family NEREID AE 

Subfamily NAMANEREINAE 

Cryptonereis n. gen. 

Small nereids that are generally similar to Namanereis but which lack frontal 
antennae. Prostomium with two pairs of eyes and two biarticulate palps. Proboscis 
without paragnaths but with a pair of toothed jaws. Peristomium with three pairs 
of tentacular cirri but not parapodia. Parapodia are sesquiramous throughout, each 
lacking a notopodial lobe but with a notopodial aciculum. Superior neuropodial 
setae are spinigerous, the rest being falcigerous. At maturity, parapodia become 
biramous with the development of capillary setae. 

TYPE SPECIES. Cryptonereis malaitae Gibbs. 



Cryptonereis malaitae n. gen. n. sp. 
Fig. 6. (A-F) 

DESCRIPTION. Larger specimens measure between 10 and 15 mm long for 55 to 
65 segments and are about 2 mm wide. 

The prostomium is rounded, roughly semi-circular in shape, with two pairs of eyes 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 



145 



and two biarticulate palps with broad palpophores. Frontal antennae are lacking 
(fig. 6. A). The peristomium carries three pairs of short tentacular cirri ; on each side 
two cirri are attached to a short base which is ventro-lateral in position and the thiid 
cirrus arises dorso-laterally on the peristomium (fig. 6. B). The proboscis lacks 
paragnaths but carries a pair of brown jaws, each with 9 or 10 teeth and a thin internal 
guard (fig. 6. c). 

The parapodia are sesquiramous in immature individuals, becoming biramous 
over most of the body when mature (see below). Notopodial lobes are lacking but 
black notopodial as well as neuropodial acicula persist (fig. 6. D). Neuropodia are 
bilobed with pointed pre-setal and rounded post-setal lips. The dorsal cirri are 
short and stout and the ventral cirri are very small. 

The neuropodial setae consist of one or two spinigers with minutely serrated 
blades (fig. 6. E) and four or five falcigers with short ciliated blades (fig. 6. F). The 
shaft and blade of the superior falciger often appear to be fused but the articulations 
of the other falcigers are usually distinct. 

Most of the specimens appear to be approaching maturity and, on preservation, 
gametes have been released from the parapodia at points just above the dorsal cirri. 




FIG. 6. Cryptonereis malaitae n.gen., n.sp. (A-B) Dorsal and lateral views of anterior 
region, (c) Left jaw in ventral view. (D) Parapodium from middle body of mature 
specimen. (E) Spiniger. (F) Falciger. 



146 P. E. GIBBS 

In these specimens, long, slender capillary setae are present in both rami of the 
parapodia from setigers 8 or 9 to the last few segments. In some, these capillary 
setae are short and few in number but in others, probably at a more advanced stage 
of maturity, they are very much longer, being about three times the length of the 
falcigers. In the latter specimens, each parapodium has a compact notopodial 
bundle, composed of about 20 capillaries, arising from between the dorsal cirrus 
and the notopodial aciculum, and a further 8 to 10 capillaries appear amongst the 
neurosetae (fig. 6. D). The specimen chosen for the holotype shows this condition. 

In the collection there is an immature individual which is only 6-5 mm long for 
35 segments (incomplete). This specimen does not possess the capillary setae found 
in the larger specimens. It is thought therefore that C, malaitae has an epitokous 
phase at maturity in which capillary (or natatory) setae (recalling those that are 
formed in the epitokes of some Syllidae) are developed and that the majority of the 
type specimens, collected on 20.xi.65, are approaching, or at, this stage of develop- 
ment. 

C. malaitae is apparently dioecious. The oocytes are yolky and those released on 
preservation by the females vary greatly in size but are mainly between 100 to 150(1 
in diameter. Spermatozoa are recognizable in the coelomic contents of the more 
mature males. In life, specimens are purplish but assume a brownish colour in 
alcohol. 

It seems inadvisable to enlarge the generic description of the relatively well-known 
and widespread genus Namanereis Chamberlin (see Hartman, 19590) to include 
those forms which lack frontal antennae, hence the need to erect a new genus. Apart 
from its lacking frontal antennae and also its possession of capillary setae at maturity, 
Cryptonereis closely resembles Namanereis and the relationship between these two 
genera parallels that between Micronereis Claparede and Micronereides Day in terms 
of the presence or absence of antennae. 

HABITAT. Between the fibres composing a leaf frond of the coconut palm found 
stranded at about MTL. 

RECORDS. Alite Harbour, Langa-Langa Lagoon (Malaita) - 14. 
British Museum (Natural History) Registration No. Holotype 1970-30 

Paratypes 1970-31 

NOTES. It is difficult to decide whether the habitat of C. malaitae is marine or, 
like many other Namanereinae, damp-terrestrial since either is possible, depending 
on the period of time the leaf frond had been immersed in seawater. Interestingly, 
further specimens of this, or a closely related species, were collected by the Expedition 
Land Party (Mr. J. Peake and Dr. J. Greenslade) in moist leaf litter at an altitude 
of about 350 m at two separate localities, namely Betimatu on Mt. Austen, Guadal- 
canal and on Kolombangara in the New Georgia Group. This material consists of 
two specimens (one incomplete and the other dried) both about 5 mm long and 
resembling immature C. malaitae in detail. However in both specimens the pros- 
tomium is damaged and because the presence or absence of frontal antennae cannot 
be determined their identity must remain in question. It is possible that these two 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 147 

specimens could be Namanereis amboinensis (Pflugfelder, 1933, as Lycastopsis 
amboinensis) which resembles the immature stage of C. malaitae in all details but 
possesses antennae. 



Namalycastis indica (Southern, 1921) 

Lycastis indica Southern, 1921 : 578, pi. 19, fig. 2. a-j, text-fig. 2. a-d. 
Namalycastis indica: De Silva, I965a : 5, fig. 2. a-e; Day, 1967 : 301, 14.2. p-s. 

HABITAT. Burrowing within the fibrous husks of Nipa palm nuts immersed in 
brackish water (5'6% ). 

RECORD. Lagoon at the mouth of the Lunga R. at Lunga Pt. - 12. 

NOTES. N. indica was found only in those Nipa nuts which were waterlogged, 
lying along and just below the water-line of the swamp. The largest specimen is 
60 mm long. 

DISTRIBUTION. Indian Ocean. 



Subfamily NEREINAE 
Ceratonereis costae (Grube, 1840) 

Ceratonereis costae : Fauvel, 1923 : 349, fig. 136. a-f; 1953, X 94 % 98. a-f; Day, 1967 : 325, fig. 
14.10. h-1. 

HABITAT. Amongst Phyllochaetopterus socialis tubes; on Halichondria sp. at 
5 m depth. 

RECORDS. Batuona Is. - i ; Yandina - i. 

DISTRIBUTION. North Atlantic; Mediterranean; Indo-west-Pacific. 



Ceratonereis erythraeensis Fauvel, 1918 

Ceratonereis erythraeensis Fauvel, 1918 : 505, fig. 2. a-k; Day, 1967 : 327, fig. 14.10. o-t. 

HABITAT. Muddy silt under boulders at MTL. 

RECORDS. Naro Bay - 20. 

DISTRIBUTION. Indian Ocean; East Australia; Japan. 



148 P. E. GIBBS 

Ceratonereis mirabilis Kinberg, 1866 

Ceratonereis mirabilis: Gravier, 1901 : 172, pi. n, fig. 42; Fauvel, 1953 : 200, fig. 103. a-c; Day, 
1967 : 324, fig. 14.10. a-g. 

HABITAT. Silty sand under boulders; in Acropora rubble; within interstices of 
Galaxea; on Halichondria sp. at 5 m depth. 

RECORDS. Tetel Is. - 2 ; Komimbo Bay - i ; Graham Pt. - 2 ; Yandina - 3. 
DISTRIBUTION. Circumtropical. 



Nereis (Neanthes) caudata (Delle Chiaje, 1825) 

Nereis cricognatha: Fauvel, 1953 : 180, fig. 91. a-c. 

Nereis (Neanthes) arenaceodonta: Pettibone, 1963 : 162, figs 44.!, 45.6. 

Nereis (Neanthes) caudata: Day, 1967 : 321, fig. 14.9. f-j. 

HABITAT. Under coral boulder in muddy silt; sand at 5 m depth. 
RECORDS. Komimbo Bay - i ; ML 229 - i. 
DISTRIBUTION. Cosmopolitan. 



Nereis (Neanthes) unifasciata Willey, 1905 

Nereis (Neanthes) unifasciata Willey, 1905 : 271, pi. 4, figs 85-88; Fauvel, 1953 : 182, 92. a-h; 
Day, 1967 : 318, fig. 14.7. u-y. 

HABITAT. Under coral boulder; in beachrock. 
RECORDS. Komimbo Bay - i ; Lauvie Is. - i. 
DISTRIBUTION. Tropical Indo-west-Pacific. 



Nereis (Neanthes) sp. cf. kerguelensis Mclntosh, 1885 

Nereis kerguelensis Mclntosh, 1885 : 225, pi. 35, figs 10-12, pi. i6A, figs 17-18. 

HABITAT. Silty sand in crevice of Porites boulder. 
RECORD. Graham Pt. - i. 

NOTES. The specimen corresponds to N. kerguelensis in all details except that the 
notopodia of the anterior segments lack intermediate pre-setal lobes, giving a bi- 
lobed, not tri-lobed structure. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 



149 



Perinereis cultrifera (Grube, 1840) 
(Fig. 7. A) 

Perinereis cultrifera : Fauvel, 1923 : 352, fig. 137. a-1; 1953 : 206, fig. 106. a-1; Day, 1967 : 337, 
fig. 14.13. o-q. 

HABITAT. Chiefly under coral boulders on sand at MTL; also in crevices in reef 
platform and in beachrock. 

RECORDS. Kokomtambu Is. -i; Tetel Is. -3; Matiu Is. -6; New Manra-32; 
Maraunibina Is. -4; Lauvie Is. -i; Graham Pt. -4; Yandina-i; Fanalei-2 
(heteronereids). 

NOTES. Two female heteronereid stages of this species are included in the 
collection made during the 'palolo' rising at Fanalei on 2.xi.66 (local name - 'falisi 
ogu'). They were taken between 2000 and 2200 hours and correspond to the 
description of this stage given by Izuka (1912) and Fauvel (1923) except that lower 
ligule of the notopodium in the modified parapodium has a small ear-shaped lobe 
arising from its ventral margin (fig. 7. A). These specimens measure 45 and 52 mm 
long for 116 and 225 segments respectively, with the modified parapodia commencing 
at setigers 20 to 22. 

DISTRIBUTION. Cosmopolitan. 





1.0mm 



1.0mm 



FIG. 7. Perinereis cultrifera (A) Modified parapodium of heteronereid (?) stage (setae 
omitted). Perinereis nuntia (B) Modified parapodium of heteronereid (<$) stage (setae 
omitted) . 



150 P. E. GIBBS 

Perinereis nigropunctata (Horst, 1889) 

Nereis (Perinereis) nigro-punctata Horst 1889 : 171, pi. 8, figs 1-3; 1924 : 171. 

Perinereis nigro-punctata: Fauvel, 1953 : 210, fig. 107. b-f; Day, 1967 : 337, fig. 14.13. r-v. 

HABITAT. Amongst Phyllochaetoptems socialis tubes; under coral boulders; 
abundant in beachrock. 

RECORDS. Batuona Is. - 2 ; Graham Pt. - 2 ; Lauvie Is. - num. 
DISTRIBUTION. Tropical Indo-west-Pacific. 

Perinereis nuntia (Savigny, 1818) 

(Fig. 7. B) 
Perinereis nuntia: Fauvel, 1953 : 212, fig. 109. a-g; Day, 1967 : 334, fig. 14.12. p-s. 

HABITAT. Chiefly found under boulders lying on clean sand and gravel at MTL. 

RECORDS. Haroro - 3 ; Tetel Is. - 3 ; Cape Esperance - 17 ; Matiu Is. - 2 ; New 
Manra - 3 ; Mouth of Lunga R. - 4 ; Yandina - 50 ; Honiara - 20 ; Vera Vera Entrance 
- i (heronereid ($} ). 

NOTES. In these specimens there are I to 3 paragnaths in group i and the parag- 
naths of group VI are usually flattened, sometimes mixed with conical forms. 
However the number of paragnaths in group V is very variable : for example, in the 
sample of 50 individuals from the Yandina population, one specimen has four 
paragnaths in group V, 26 have three arranged in a triangle, n have two and 12 
have only one. Thus in terms of the subspecific forms (see Fauvel, 1953) Perinereis 
nuntia brevicirrus composes about half the population and P. nuntia vallata about a 
quarter, the remaining quarter being intermediate in character between these two 
forms. 

The male heteronereid stage of P. nuntia brevicirrus was captured swimming near 
the surface at Vera Vera Entrance on 8.xi.65 at 1500 hours. It measures 90 mm 
long for about 150 segments, the modification of the parapodia starting between 
setigers 23 to 27. The structural details of the modified parapodium (fig. 7. B) follow 
those described and figured by Izuka (1912, as N. mictodonta). 

DISTRIBUTION. South Africa; Indo-Pacific. 



Platynereis insolita Gravier, 1901 

Platynereis insolita Gravier 1901 : 197, pi. 12, fig. 53, text-figs 203-206; Day, 1967 : 307, fig. 
14.4.1. 

HABITAT. Under coral boulder; amongst Phyllochaetopterus socialis tubes and 
Amphiroa growth. 

RECORDS. Matiu Is. - i ; Komimbo Bay - i ; New Manra - 2. 
DISTRIBUTION. Indian Ocean. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 151 

Pseudonereis anomala Gravier, 1901 

Pseudonereis anomala Gravier, 1901 : 191, pi. 12, figs 50-52, text-figs 194-202; Fauvel, 1953 : 
217, fig. no. e-g; Day, 1967 : 333, fig. 14.12. g-j. 

HABITAT. Abundant amongst Phyllochaetopterus socialis tubes and Amphiroa; 
burrowing in reef platform and beachrock. 

RECORDS. Mamara Pt. - i ; Batuona Is. - num. ; New Manra - num. ; Yandina - 
6 ; Maramasike Pg. - 3. 

DISTRIBUTION. Indo-Pacinc. 



Pseudonereis masalacensis (Grube, 1878) 

Nereis (Lycoris) masalacensis Grube, 1878 : 75, pi. 5, fig. 4. 
Pseudonereis masalacensis: Fauvel, 1947 : 51, fig. 49. a-k. 

HABITAT. Under coral boulders on sand at MTL; amongst Phyllochaetopterus 
socialis tubes and algal growth (Amphiroa). 

RECORDS. Batuona Is. - 2 ; New Manra - 3 ; Yandina - 2. 
DISTRIBUTION. West and South Pacific. 



Pseudonereis variegata (Grube, 1857) 

Pseudonereis gallapagensis : Fauvel, 1953 : 2I 5. fig- IIQ - a ~ c - 
Pseudonereis variegata: Day, 1967 : 331, fig. 14.12. a-f. 

HABITAT. Burrowing in reef platform and beachrock; amongst Amphiroa; under 
boulders on clean sand. 

RECORDS. Matiu Is. - 8 ; Sifola - 8 ; New Manra - 2 ; Lauvie Is. - 3 ; Yandina - 2. 
DISTRIBUTION. Circumtropical. 



Solomononereis n. gen. 

Prostomium with two antennae, two biarticulate palps and two pairs of eyes. 
Peristomium with four pairs of tentacular cirri, dorsal pairs long, ventral pairs much 
shorter. Proboscis with small, rod-like chitinous paragnaths arranged in eight 
discrete groups on the maxillary ring; oral ring without paragnaths. Parapodia 
biramous with spinigerous setae in both rami throughout the body; falcigerous setae 
in both rami in the middle and posterior segments. 

TYPE SPECIES. Solomononereis marauensis Gibbs 



152 



P. E. GIBBS 



Solomononereis marauensis n. gen. n. sp. 



(Fig. 8. A-H) 

DESCRIPTION. The holotype measures 45 mm for 123 segments and is incomplete 
posteriorly. The largest of the three paratypes is 40 mm for 157 segments. The 
width of the body is about 3 mm. 

The prostomium is rectangular in shape with two short, widely spaced antennae, 
two pairs of eyes, and a pair of palps with stout palpophores, often with a constriction 
just below the tip, and small palpostyles (fig. 8. A). The peristomium is weakly 
developed and is present as a narrow band on the dorsal surface but does not form a 
distinct ring ventrally. Of the four pairs of tentacular cirri, the two dorsal pairs 
are very long, extending back as far as setiger 20 but the ventral pairs are short, 
reaching to setiger 3 only. 




FIG. 8. Solomononereis marauensis n.gen., n.sp. (A) Dorsal view of anterior region and 
proboscis. (B) Ventral view of proboscis, (c) Paragnaths of Group II. (D-F) Anterior, 
middle and posterior parapodia. (G) Notopodial falciger. (H) Subacicular neuro- 
podial spiniger. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 153 

The proboscis carries a pair of brown, curved jaws, each with 7 or 8 teeth. 
The chitinous paragnaths are confined to the maxillary ring ; they are minute, rod-like 
(7-iO{/. in diameter) structures which are closely packed together in eight groups, 
three on the dorsal side and five on the ventral (fig. 8. A, B). Each of these discrete 
groups is composed of 10 to 15 paragnaths (fig. 8. c). 

The structure of the parapodia changes progressively throughout the body and 
those of the anterior region are larger than those of the middle and posterior parts of 
the body. In the anterior region, the notopodium has two pointed ligules and the 
neuropodium has a bilobed upper ligule, composed of a pointed pre-setal lip and a 
rounded post-setal lip, and a pointed lower ligule (fig. 8. D). Between setigers 20 and 
30, the upper notopodial ligule and the lower neuropodial ligule decrease in size, 
the former being lost altogether in more posterior segments while the latter persists 
as a rudiment (fig. 8. E, F). The dorsal cirri are slightly longer than the parapodial 
lobes in anterior segments but are relatively much longer in the posterior segments 
because of the diminished size of the parapodia. 

At the base of the dorsal cirri, there are conspicuous glands which are roughly 
oval in shape and have a yellowish colour in preserved (alcohol) specimens. Each 
gland consists of fascicles of spindle-shaped cells which appear to open to the exterior 
at a pore situated just above the base of the dorsal cirrus. 

Spinigerous setae are present in both notopodial and neuropodial bundles through- 
out the body. The notosetae are all homogomphic while the neurosetae vary from 
homogomphic to hemigomphic in the superior position, becoming heterogomphic 
in the inferior part. Superior sub-acicular spinigers have coarsely serrated blades 
(fig. 8. H) but all others are finely serrated. In the region of setiger 40, and persisting 
to the posterior end, two or three homogomph falcigers (fig. 8. G) appear in the 
notopodial bundle and, in addition, a single heterogomph falciger appears in the 
supra-acicular part of the neuropodium. Each falciger has an elongate, ciliated 
blade with a curved tip supported by a distinct tendon. All setae have a crystalline 
appearance. 

A dark brown pigment remains over the dorsal surface of the proboscis, prostomium 
and anterior segments in preserved specimens. 

Solomononereis resembles Ceratonereis in having paragnaths only on the maxillary 
ring of the proboscis but this similarity is superficial, the shape, as well as the arrange- 
ment, of the 'toothpick' or rod-shaped paragnaths possessed by Solomononereis 
being very different to that of the conical paragnaths found in Ceratonereis. Other 
features separating Solomononereis from other nereid genera include the weakly 
developed peristomial ring and the arrangement of the setae, i.e. spinigers are 
present in both rami throughout the body and, in addition, a few falcigers appear in 
both rami of the middle and posterior parapodia. 

HABITAT. In sticky mud above MTL close to freshwater outflows. 

RECORDS. Fintry Pt. - i (holotype) ; Nggela Is. (Sandfly Passage) - 3. 
British Museum (Natural History) Registration No. Holotype 1970-32 

Paratypes 1970-33 



154 p - E - GIBBS 

Family NEPHTYIDAE 
Nephtys (Aglaophamus) munamaorii n. sp. 

(Fig. 9. A-B) 

DESCRIPTION : The holotype (from ML 37) is the largest specimen in the col- 
lection. It measures 16 mm in length for 60 segments and is about i mm wide. 

The prostomium is rectangular in shape, its length being about i -5 times the width, 
and has four small antennae on the antero-lateral margins. The proboscis has a 
long median papilla on its dorsal side with 14 rather indistinctly aligned rows of 
5 or 6 papillae. Two small eyes, which appear as black spots, are situated near the 
posterior border of the second segment. 

Branchiae commence on setiger 3 and are well developed in the anterior half of the 
body but gradually diminish in size in the posterior half so as to be absent from the 
last 10 or 12 segments. Each branchia is involute with a dorsal cirrus at its base 
(fig. 9. A). 

The setigerous lobes of the parapodial rami are pointed. In the anterior segments, 
both the pre-setal and the post-setal lamellae are rounded, the latter being slightly 
larger, but in posterior segments these lamellae are much reduced in size (fig. 9. B). 
There are no superior neuropodial lamellae. 

Setae are of the three usual types. Barred (or laddered) -capillary setae compose 
the anterior fan which also includes a few smooth capillaries : in posterior segments 
the barred setae are longer and more slender with less distinct bars. Smooth or 
minutely serrated capillaries form the posterior fan : in the latter, there is also a row 
of lyriform forked setae. 

Four species belonging to the subgenus Aglaophamus have been described from the 
tropical Indo-west-Pacific region. These are N. (A.) dibranchis Grube, N. (A.) 





0.5mm 



B 



FIG. 9. Nephtys (Aglaophamus) munamaorii n.sp. (A-B) Anterior views of parapodia 
from setigers 19 and 45 respectively (setae omitted). 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 155 

lyrochaeta Fauvel, N. (A.) mirasetis Hoagland, and N. (A.) sinensis Fauvel. In all 
four species the neuropodia of anterior segments have superior lamellae. The 
absence of this lamella, in addition to further differences in the parapodial structure, 
particularly the shape of the pre- and post-setal lamellae, is sufficient to distinguish 
N. (A.) munamaorii. 

HABITAT. Mud and silty sand, 2-22 m depth. 

RECORDS. ML 37-2 (including holotype) ; ML 69 - i ; ML 157 - i ; ML 188 - 8; 
ML 194 - 2 ; ML 196 - i ; ML 218 - 2. 
British Museum (Natural History) Registration No. Holotype 1970-34 

Paratypes 1970-35-41 



Nephtys (Micronephthys) sphaerocirrata Wesenberg-Lund, 1949 

Nephthys sphaerocirrata Wesenberg-Lund, 1949 : 294, figs 24-26. 
Nephtys (Micronephthys) sphaerocirrata: Day, 1967 : 347, fig. 15.3. a-d. 

HABITAT. Silty sand at 15 m depth. 

RECORD. ML 116 - i. 

NOTES. The single specimen is an anterior fragment of 33 segments, 5 mm in 
length. It differs from the type specimen in having two pairs of reddish eyes situated 
near the posterior margin of the prostomium. These may represent a sexual 
character since the specimen is a mature female containing oocytes. Other details 
agree with the type description. 

DISTRIBUTION. Persian Gulf; South Africa; Marshall Islands. 



Nephtys (Nephtys) sp. cf. palatii Gravier, 1904 
Nephthys palatii Gravier, 1904 : 472; 1906 : 129, pi. i, figs 163-164, text-figs 286-289. 

HABITAT. A wide variety of deposits from mud to coarse sand, 2-18 m depth. 

RECORDS. ML 39 - i ; ML 68 - i ; ML 97 - i ; ML 156- i ; ML 191 - i ; ML 192 - 1 ; 
ML 196 - i ; ML 203 - i ; ML 229 - 2. 

NOTE. These specimens are close to N. palatii but the identity is uncertain 
because, on dissection, a median dorsal papilla could not be seen on the proboscis. 



Family LACYDONIIDAE 
Paralacydonia weberi Horst, 1923 

Paralacydonia weberi Horst, 1923 : 221, figs 1-2; Fauvel, 1953 : 129, figs 65. e-f. 
HABITAT. Mud and sand, 2-24 m depth. 



156 P. E. GIBBS 

RECORDS. ML 29 - i ; ML 190 - i ; ML 191 - i ; ML 230 - i. 

NOTE. The largest of the specimens is 10 mm long for 50 segments and is much 
smaller than the holotype (40 mm for 100 segments) which was dredged from deep 
water (959 m). 

DISTRIBUTION. Burma; East Indies; Samoa; Bass Strait. 



Family GLYCERIDAE 

Subfamily GLYCERINAE 

Glycera gigantea Quatrefages, 1865 

Glycera gigantea : Fauvel, 1923 : 387, fig. 152. d-k; 1953 : 296, fig. 152. d-k; Day, 1967 : 362, 
fig. 16.2. 1-n. 

HABITAT. A wide variety of deposits, chiefly silty mud and sand, from above 
MTL to 33 m depth. 

RECORDS. Tetel Is. -i; Florida Is. -3; Komimbo Bay -7; New Manra-i; 
Graham Pt. -8; Fintry Pt. -4; Maraunibina Is. -i; ML 229-1. (ML stations 
100, no, 192, 195 - one juvenile (?) at each). 

DISTRIBUTION. North Atlantic; Mediterranean; Laccadive Sea; New Britain; 
N.E. Australia; Japan; eastern North Pacific. 



Glycera lancadivae Schmarda, 1861 

Glycera lancadivae Schmarda, 1861 : 95, with text-figs; Fauvel, 1953 : 291, fig. 147. g-h; Day, 
1967 : 359- 

HABITAT. A wide variety of deposits, from fine coral mud to coarse sand and 
shell gravel from above MTL to n m depth. 

RECORDS. Tetel Is. -3; Komimbo Bay -4; Naro Bay-i; Matiu Is. -11; 
Maraunibina Is. - 22 ; Graham Pt. - 8 ; Fintry Pt. - 9 ; Yandina - 4; ML 98 - i ; 
ML 203 - i ; ML 229 - 2. 

DISTRIBUTION. Tropical Indian Ocean; N.E. Australia; New Caledonia. 



Glycera longipinnis Grube, 1878 

Glycera longipinnis Grube, 1878 : 182, pi. 8, fig. 9; Fauvel, 1932 : 125, pi. 4, figs 11-14; I 953 : 
291, fig. 148. a-d; Day, 1967 : 356, fig. 16.1. a-f. 

HABITAT. Muddy silt (Thalassia flat) at LWM. 
RECORD. Komimbo Bay - i. 
DISTRIBUTION. Tropical Indo-west-Pacific. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 157 

Glycera rouxi Audouin & Milne-Edwards, 1833 

Glycera rouxi: Fauvel, 1923 : 389, fig. 153. a-c; 1953 : 2 97. fig- J 49- a -d; Day, 1967 : 362, fig. 
16.3. a-d. 

HABITAT. Silty mud at 9 m depth. 
RECORD. ML 38-1. 

DISTRIBUTION. North Atlantic; Mediterranean; Indo-west-Pacific to Japan; 
California. 



Family EUNICIDAE 
Subfamily EUNIGINAE 
Eunice afra Peters, 1854 

Eunice afra: Grassland, 1904 : 289, pi. 20, fig. 1-5, text-figs 43-45; Fauvel, 1953 : 2 35> fig- IJ 6. 
h-i; Day, 1967 : 392, fig. 17.5. a-e. 

HABITAT. Boring in coral on reef platform and in beachrock; amongst Phyl- 
lochaetopterus socialis tubes at LWM. 

RECORDS. Matiu Is. -4; Mamara Pt. -2; Batuona Is. -2; New Manra-i; 
Lauvie Is. - 2; Lingatu - 4; Yandina - i ; Maramasike Pg. - 5. 

DISTRIBUTION. Indo-west-Pacific. 



Eunice antennata (Savigny, 1820) 

Eunice antennata: Crossland, 1904 : 312, pi. 22, figs 1-7, text-figs 56-60; Fauvel,! 953 : 240, 
fig. 118. f-g; Day, 1967 : 384, fig. 17.2. k-q. 

HABITAT. Boring in coral and under coral boulders on reef platform. 
RECORDS. Matiu Is. - 5 ; Graham Pt. - 2 ; Yandina - i. 
DISTRIBUTION. North Atlantic; tropical Indo-Pacific. 



Eunice aphroditois (Pallas, 1788) 
Eunice aphroditois: Fauvel, 1953 : 233, fig. 117. a-g; Day, 1967 : 389, fig. 17.4. l-o. 

HABITAT. In Acropora rubble and under coral boulder on reef platform at LWM. 
RECORDS. Tetel Is. - i ; Maraunibina Is. - 2. 

DISTRIBUTION. North Atlantic; Mediterranean; Indo-Pacific to Japan and 
southern California. 



158 P. E. GIBBS 

Eunice coccinea Grube, 1878 

Eunice coccinea Grube, 1878 : 153, pi. 9, fig. i; Grassland, 1904 : 297, pi. 20, figs 6-7, text-figs 
46-51; Fauvel, 1953 : 2 36, fig- 118. a-e; Day, 1967 : 389. 

HABITAT. Boring in coral on reef platform. 

RECORDS. Komimbo Bay - i ; Mamara Pt. - i ; New Manra - i ; Lingatu - i ; 
Maramasike Pg. - 2; Fanalei - 2 (posterior fragments). 

NOTES. The two posterior fragments in the collection from the Fanalei 'palolo' 
rising (2-3.ix.66) can be identified by their relative size, absence of gills, parapodial 
structure and setae. One is female and contains large spherical oocytes (360(0. in 
diameter). On one side of each of these oocytes there is a patch of dark green 
pigment, surrounding a clear circular spot. The time of spawning is recorded as 
midnight to 0600 hours. The local name is 'raka-raka'. 

DISTRIBUTION. Gulf of Guinea; tropical Indo-west-Pacific. 



Eunice grubei Gravier, 1900 

Eunice grubei Gravier, 1900 : 258, pi. 14, figs. 87-88, text-figs 125-129; Fauvel, 1953 : 237, fig. 
119. a-e; Day, 1967 : 391. 

HABITAT. Sedimented crevices in Forties boulders. 
RECORDS. Graham Pt. - 2. 

NOTES. One specimen is complete and, in this, the branchiae, which are present 
only on the anterior half of the body, have a maximum of 4 or 5 filaments. 

DISTRIBUTION. Tropical Indo-west-Pacific. 



Eunice marovoi n. sp. 

(Fig. 10. A-H) 
Eunice sp.: Gibbs, 1969 : 455, fig. 133. 

DESCRIPTION. Complete specimens from the dredged material have a maximum 
length of 15 mm for 60 to 70 segments and a width of i-o to 1-5 mm. Larger 
specimens discovered living on Cerithium vertagus are all incomplete but are 
estimated to have been 20 to 25 mm in length. The holotype from ML 96 is 15 mm 
long for 66 segments. 

The prostomium is bilobed with a shallow median groove between the palps 
(fig. 10. A). The antennae are weakly annulated and the median is about twice 
as long as the laterals, which, in turn, are about equal in length to the tentacular cirri 
(fig. 10. B). The eyes are black and reniform in shape. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 



159 



The mandibles are elongate calcified plates. The maxillae are almost transparent 
but the toothed edges are brownish in colour and the medial and anterior edges of 
the maxillae supports ("carriers") are black, forming a T-shaped figure (fig. 10. c). 
Due to the transparent nature ot the maxillae the details are difficult to determine. 
Maxillae I are falcate and devoid of basal teeth. The left Maxilla II is falcate with 
4 to 6 ill-defined basal teeth. The right Maxilla II, left Maxilla III and Maxillae IV 
all have 10 to 13 small teeth while Maxillae V are toothless plates. The maxillary 
formula thus reads: MX. 1 = 1 + 1; MX. II = (i + (4 6)) + (10 13) ; MX. Ill 
= (10 13) + o; MX. IV = (10 13) + (10 13). 

Branchiae commence on setiger 3 on all specimens and these are strongly arched 
over the dorsum, with a maximum of 5 or 6 filaments (fig. 10. D). In the larger 
specimens there are 21 to 23 pairs (i.e. extending to setigers 23 to 25) but smaller 




FIG. 10. Eunice marovoi n.sp. (A) Dorsal view of prostomium. (B) Antero-lateral view 
of anterior region, (c) Jaws. (D-E) Branchia and parapodium of setiger 10. (F) Comb 
seta. (G) Falciger. (H) Acicular seta. 



160 P. E. GIBBS 

specimens have only 15 to 20 pairs. The dorsal cirri are shorter but slightly stouter 
than the branchial filaments. On branchiferous segments, the ventral cirri are 
short and are swollen at their bases (fig. 10. E), a feature that is absent in the more 
posterior segments. 

Acicula and setae are transparent. The capillary setae are slender with finely 
serrated edges. Comb setae are few in number and have about 7 teeth, one of the 
outer pair being greatly elongated (fig. 10. F). The falcigerous setae have bidentate 
blades with faintly serrated guards (fig. 10. G). Acicular setae are tridentate with 
rounded guards (fig. 10. H) ; two or three occur in each parapodium from setigers 15 
to 20 to the posterior end. The projecting tips of the acicula are rounded and slightly 
curved. 

The colour of preserved specimens is white. Despite their small size, a number of 
the specimens are mature with coelomic gametes. It would appear therefore that 
E. marovoi does not exceed 20 to 25 mm in length even when fully grown. 

E. marovoi is distinct from all other Eunice species known from the Indo-west- 
Pacific region in terms of its maxillary structure, the number of branchiae and 
branchial filaments, in combination with the structure of the setae. 

HABITAT. A wide variety of deposits ranging from mud to coarse sand from 
LWM to 33 m depth. At LWM on a sand flat to the north of Tankomo Is. (in the 
southern part of Marovo Lagoon) E. marovoi was discovered attached to the shell 
of the gastropod Cerithium vertagus L. 

RECORDS. ML 37 - i ; ML 96-1 (holotype) ; ML 97 - i ; ML 100 - i ; ML no - 9 ; 
ML 134 - i ; ML 192 - 2 ; ML 218 - i ; ML 229 - i ; ML 230 - 2 ; north of Tankomo 

Is. -5. 

British Museum (Natural History) Registration No. Holotype 1970-42 

Paratypes 1970-43-52 

NOTES. The tube of this species is constructed of coarse bottom material, such 
as scaphopod and bivalve she)l fragments, cemented to an organic lining. In those 
specimens associated with Cerithium vertagus, the tube was constructed along the 
length of the upper surface of the gastropod shell (see Gibbs, 1969). 



Eunice norvegica (Linnaeus, 1767) 

Eunice floridana : Fauvel, 1923 : 402, fig. 157. a-g; 1953 : 235, fig. 117. a-g. 
Eunice norvegica: Pettibone, 1963 : 240, fig. 63. f; Day, 1967 : 388, fig. 17.3. r-v. 

HABITAT. On surface of Acropora sp. 

RECORD. Yandina - 1. 

NOTES. The specimen is 17 mm long, 3 mm wide and has 70 segments (apparently 
regenerating from setiger 27). Its preserved colour is medium brown with three 
whitish spots per segment arranged in longitudinal lines down the body; the gills 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 161 

and dorsal cirri are purple. In life the colour was bluish-black with orange spots. 
The maxillary formula is: MX. I = i -|- i; MX. II = 7 -j- 7; MX. Ill = 6 + o; 
MX. IV = 4 + 8; MX. V = i + i. The labrum is calcined and pearly- white in 
colour. The branchiae start on setiger 5 and have a maximum of 6 or 7 filaments. 
Acicular setae are black with conspicuous guards. 

In specimens from other localities the branchiae commence between setigers 7 to 
10 but otherwise this specimen agrees in detail. Apart from a single specimen 
from Indo-China (Fauvel, 1939) no other records of E. norvegica from the Pacific 
Ocean have been traced despite the species being widespread and occurring in depths 
of over 1500 m (Pettibone, 1963). 

DISTRIBUTION. North Atlantic ; Mediterranean ; Indian Ocean ; Indo-China. 



Eunice tentaculata Quatrefages, 1865 

Eunice tentaculata: Fauvel, 1953: 234, fig. 118. m-p; Day, 1967 : 391, fig. 17.4. s-v. 

HABITAT. Under coral boulders on silty sand ; amongst Phyllochaetoptems socialis 
tubes. 

RECORDS. Maraunibina Is. - 2; Batuona Is. - i. 
DISTRIBUTION. Indo-west-Pacific. 



Eunice tubifex Grassland, 1904 

Eunice tubifex Grassland, 1904 : 303, pi. 21, figs 1-8, text-figs 52-55; Fauvel, 1953 : 2 3 2 n g- Il( >. 
a-g; Day, 1967 : 386, fig. 17.3. k-q. 

HABITAT. In coral at LWM and on wharf piles at 5 m depth. 
RECORDS. Tetel Is. - 2 ; Yandina - 2. 
DISTRIBUTION. Tropical Indo-west-Pacific. 



Eunice (Palola) siciliensis Grube, 1840 

Eunice (Palola} siciliensis: Fauvel, 1923 : 405, fig. 159. e-m; 1953 : 241, fig. 121. e-m; Day, 1967 : 
382, fig. 17.2. a-f. 

HABITAT. Boring in coral and beachrock; in Acropora rubble and shell gravel; 
under coral boulders in silty sand. 

RECORDS. Kokomtambu Is. - i ; Tetel Is. - 16 ; Komimbo Bay - 2 ; Mamara Pt. - 
3 ; Matiu Is. - 34 ; Yandina - 2 ; Lingatu - 6 ; New Manra - 9 ; Lauvie Is. - n ; 
Graham Pt. - 4; Maramasike Pg. - 10; Fanalei - 4 (posterior fragments). 



162 P. E. GIBBS 

NOTES. The four posterior fragments collected during the spawning at Fanalei 
can be readily identified by the absence of comb and acicular setae, the black 
acicula and the dark spot on the ventral surface of each segment. Three of the 
fragments are female and contain spherical oocytes about 2OO[A in diameter. The 
time of spawning on 2.xi.66 was between 2000 and 2200 hours. The local name is 
"falisi ogu". 

DISTRIBUTION. Mediterranean; circumtropical. 



Lysidice collaris Grube, 1870 
Lysidice collaris: Fauvel, 1953 : 248, fig. 124. a-g; Day, 1967 : 402, fig. 17.8. a-f. 

HABITAT. Boring in coral and beachrock; amongst Phyllochaetopterus socialis 
tubes ; in the sponge Halichondria sp. at 5 m depth. 

RECORDS. Matiu Is. - 5 ; Batuona Is. - 1 ; New Manra - 3 ; Sifola - 2 ; Lingatu - 2 ; 
Mamara Pt. - 2 ; Graham Pt. - i ; Lauvie Is. - i ; Yandina - 12 ; Fanalei - i. 

NOTES. The specimen from the Fanalei "palolo" rising is spent. It has enlarged 
eyes provided with lenses. 

DISTRIBUTION. Tropical Indo-Pacific to Japan and Gambier Islands. 



Marphysa macintoshi Crossland, 1903 

Marphysa macintoshi Crossland, 1903 : 137, pi. 14, figs 3-6, text-fig. 12; Fauvel, 1953 : 246; 
Day, 1967 : 396, fig. 17.6. a-e. 

HABITAT. Silty mud at MTL. 
RECORDS. Nggela Is. - 3. 
DISTRIBUTION. Tropical Indian Ocean. 



Nematonereis unicornis (Grube, 1840) 

Nematonereis unicornis: Fauvel, 1923 : 412, fig. 162. h-n; 1953, 249, fig. 124. h-n; Day, 1967 : 403, 
fig. 17.8. j-n. 

HABITAT. Under boulders in muddy silt at MTL; amongst Phyllochaetopterus 
socialis tubes at LWM. 

RECORDS. Tetel Is. - i ; Batuona Is. - i. 

DISTRIBUTION. North Atlantic; Mediterranean; tropical Indo-west-Pacific. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 163 

Subfamily ONUPHINAE 
Onuphis (Nothria) holobranchiata Marenzeller, 1879 

Onuphis (Nothria] holobranchiata Marenzeller, 1879 : 132, pi. 4, fig. i; Izuka, 1912 : 106, pi. n, 
figs 10-12; Fauvel, 1953 : 2 56, fig- 127. f-h; Day, 1967 : 424, fig. 17.13. f-g. 

HABITAT. A wide variety of deposits, from silty sand to coarse sand and shell 
gravel from MTL to LWM ; mud with organic debris at 2 m depth. 

RECORDS. Tetel Is. -14; Komimbo Bay-i; Matiu Is. -i; Kalota Is. -i; 
ML 68 - 37. 

NOTES. The distribution of the simple branchiae over the anterior segments 
shows some degree of variation, perhaps reflecting differences in the growth stages. 
For example, in the smaller specimens present in the large sample obtained at ML 68 
the branchiae commence on setigers 2 to 5 whereas in the larger specimens from the 
same sample they begin on the first setiger, as typical for the species. Specimens 
of all sizes possess similar pseudocompound hooks. 

The dredged specimens were chiefly extracted from tubes composed of plant 
debris and sediment particles cemented to an organic lining and often the tubes were 
found to run along the central cavities of hollow plant stems. Specimens from the 
littoral zone had tubes encrusted with coarse sand grains and Halimeda discs. 

DISTRIBUTION. Tropical Indo-west-Pacific to Japan. 



Subfamily LYSARETINAE 
Oenone fulgida (Savigny, 1818) 

Aglaurides fulgida: Fauvel, 1953, 250, fig. 125. a-f. 

Oenone fulgida: Crossland, 1924 : 85, fig. 106-111; Imajima & Hartman, 1964 : 267; Day, 
1967 : 426, fig. 17.14. a-g. 

HABITAT. Boring in coral (Porites). 
RECORDS. Tetel Is. - 2 ; Yandina - i. 
DISTRIBUTION. Circumtropical. 



Subfamily LUMBRINERINAE 
Lumbrineris latreilli Audouin & Milne-Edwards, 1834 

Lumbriconereis latreilli: Fauvel, 1923 : 431, fig. 171. m-r; 1953 : 266, fig. 134. m-r. 
Lumbrineris latreilli: Day, 1967 : 438, fig. 17.16. p-t. 

HABITAT. Under coral boulders in silty sand; mud at 18 m depth. 



164 P. E. GIBBS 

RECORDS. Tetel Is. - I ; Komimbo Bay - i ; Graham Pt. - 2 ; Maraunibina Is. - I ; 
ML 188 - i. 

DISTRIBUTION. Cosmopolitan. 



Lumbrineris papillifera (Fauvel, 1919) 

Lumbriconereis papillifera Fauvel, igiga, : 395, pi. 15, figs 9-16. 
Lumbrineris papillifera: Day, 1967 : 442, fig. 17.17. p-s. 

HABITAT. Silty sand at 9 m depth. 
RECORD. ML 56-1. 

NOTES. The specimen is incomplete posteriorly but middle segments having the 
characteristic ventral papillae are present and other details correspond to the species 
description. This is the first record of the species from the Pacific region. 

DISTRIBUTION. Tropical East Africa. 



Lumbrineris sphaerocephala (Schmarda, 1861) 

Notocirrus sphaerocephalus Schmarda, 1861 : 116, with text-figs. 

Lumbriconereis sphaerocephala: Ehlers, 1904 : 33, pi. 5, figs 3-11; Fauvel, 1953 : 267, fig. 135. c-f. 

HABITAT. Clean sand at LWM. 
RECORD. Komimbo Bay - i. 
DISTRIBUTION. Indo-west-Pacific. 



Subfamily ARABELLINAE 
Arabella tricolor (Montagu, 1804) 

Arabella iricolor: Fauvel, 1923 : 438, fig. 175. a-h; 1953 : 2 74 ^g- I 4- a ~^'> Day, 1967 : 446, 
fig. 17.18. i-m. 

HABITAT. Under boulders in muddy silt, sand and shell gravel at MTL. 
RECORDS. Tetel Is. - 3; Maraunibina Is. - 2; Graham Pt. - 6. 
DISTRIBUTION. Cosmopolitan. 



Arabella mutatis (Chamberlin, 1919) 

Arabella mutans: Fauvel, 1953 : 275, figs 140. i-1, 143. g-i; Day, 1967 : 446, fig. 17.18. f-h. 
HABITAT. Amongst Phyllochaetopterus socialis tubes at LWM. 
RECORDS. Batuona Is. - 5. 
DISTRIBUTION. North Atlantic; East Africa; Easter Is.; eastern Pacific. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 165 

Drilonereis major Grassland, 1924 

Drilonereis major Crossland, 1924 : 57, figs 73-79; Fauvel, 1953 : 277, fig. 143. k-1. 

HABITAT. Boring (?) in coral on reef platform at LWM. 
RECORDS. Maramasike Passage - 2. 

NOTES. The prostomium is semi-circular and flattened. The maxillary formula 
is MX. I = i + i ; MX. II = 6 + 6; MX. Ill = (3 4) + (3 4); MX. IV = (2 3) 
+ (2 3) : the teeth of MX. Ill and MX. IV are very irregular in size and difficult 
to count, as noted by Fauvel (1932). This is a new record for the Pacific region. 

DISTRIBUTION. Gulf of Suez; Zanzibar; Bay of Bengal. 



Subfamily DORVEILLEINAE 
Dorvillea sp. 

HABITAT. Coarse sand at MTL. 
RECORD. Komimbo Bay - i. 

NOTES. The specimen is 7 mm long (incomplete). Its antennae have eight 
annuli and the dorsal cirri, commencing on setiger 2, are mounted on long cirrophores. 
Forked setae are present from the first setiger and have unequal prongs. It is 
close to D. incertus (Schmarda) but differs from this species in having forked setae 
from the first, not the second, setiger. 



Family ORBINIIDAE 
Haploscoloplos bifurcatus Hartman, 1957 

Haploscoloplos bifurcatus Hartman, 1957 : 2 77- 

HABITAT. Mud and silty sand, 2-13 m depth. 
RECORDS. ML 68 - 16 ; ML 69 - i ; ML 228 - 2. 

NOTES. The thorax is composed of 17 to 20 setigers. Branchiae commence on 
setigers 8, 9 or 10 as small triangular lobes, increasing in size on the posterior thoracic 
segments. Characteristically, the post-setal lobes of the neuropodia in the posterior 
half of the thorax (from setiger 12) are divided. Abdominal parapodia lack inter- 
ramal cirri. 

DISTRIBUTION. South and south-east Australia. 



166 P. E. GIBBS 

Naineris laevigata (Grube, 1855) 

Naineris laevigata: Fauvel, 1927 : 22, fig. 7. a-1; 1953 : 310, fig. 163. a-1; Hartman, 1957 : 2 97. 
pi. 35, figs 1-8; Day, 1967 : 539, fig. 32.2. a-f. 

HABITAT. Medium to coarse sand, MTL to LWM. 
RECORDS. Haroro - i ; Graham Pt. - 2. 
DISTRIBUTION. Cosmopolitan in warm waters. 



Family SPIONIDAE 
Dispio maroroi n. sp. 

(Fig. ii. A-D) 

DESCRIPTION. All four specimens are incomplete. The holotype is 18 mm in 
length with 46 segments and the largest of the paratypes is 45 mm long for 86 seg- 
ments. The diameter of the body is about 2 mm. 

The prostomium is pointed anteriorly and extends back as an occipital ridge to 
the first segment where it forms a small occipital tentacle (fig. n. A). On either 
side of this ridge there are conspicuous nuchal slits which are concealed at their 
posterior ends by dorsal extensions of the peristomium. Two small eyes, not 
visible from the dorsal aspect, are situated laterally on the prostomium. The palps 
are missing on all specimens. 

The parapodia of the first two segments differ from those of subsequent segments 
in that the post-setal lobes of the notopodia are expanded, with 6 or 7 digitiform 
processes along their free edges (fig. n. B). The lobes of subsequent notopodia have 
smooth outlines. The post-setal neuropodial lobes of the first few segments are 
rounded, becoming flatter in shape in later segments. Pre-setal lobes are low and 
rounded. 

Branchiae are present on all segments from setiger i. They are fused with the 
post-setal notopodial lobes for most of their length and are slightly longer than the 
lobes. Commencing between setigers 18 to 20, accessory branchiae appear on the 
posterior side of the notopodia. Each is a palmate group of 6 to 8 lobes, each lobe 
containing a vascular loop (fig. 11. c). 

Apart from the first setiger, which has a bundle of long fine capillaries which 
projects forward over the head region, the notosetae are broad- winged and narrow- 
winged capillaries, the two types being of similar length. The neurosetae of the 
anterior segments are similar to the notosetae except that the narrow-winged 
capillaries are longer than the broad-winged. Between setigers 24 to 28, the broad- 
winged capillaries are replaced by very finely pointed setae and 4 or 5 unidentate 
hooded hooks appear (fig. n. D). In all neuropodia, there is an inferior group of 
4 or 5 sabre-like setae. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 



167 



D. mar or oi is close to the type species D. uncinata Hartman (1951) from Florida 
but differs from it in that the post-setal notopodial lobes of setigers I and 2 are much 
enlarged whilst that of setiger 3 is unmodified. A further difference is that the 
accessory branchiae start between setigers 18 to 20 not between setigers 24 to 28 and 
also there are minor variations in the form of the head region. In other details the 
specimens from the Solomon Islands resemble those from Florida. The central 
American species D. remanei and D. schusterae, described by Friedrich (1956), both 
differ in having digitate neuropodial, as well as notopodial, lobes in the anterior 
segments. In the fourth species, D. magna (Day, 1955) the accessory branchiae 
take the form of minute lamellae, not lobes. 

HABITAT. Clean sand at LWM. 

RECORDS. Komimbo Bay - i (holotype) ; Graham Pt. - 3. 
British Museum (Natural History) Registration No. Holotype 1970-53 

Paratypes 1970-54 





FIG. n. Dispio maroroi n.sp. (A) Dorsal view of head region. (B-C) Posterior views of 
parapodia from setiger i (setae omitted) and setiger 40. (D) Neuropodial hook. 



Laonice cirrata: Fauvel, 1927 
fig. 18.6. h-k. 



Laonice cirrata (Sars, 1851) 
: 38, fig. 12. a-e; 1953 : 315, i 



HABITAT. Mud at n m depth. 
RECORD. ML 195 - i. 
DISTRIBUTION. Cosmopolitan. 



j. 165. a-e; Day, 1967 : 480, 



168 P. E. GIBBS 

Malacoceros indicus (Fauvel, 1928) 

Scolelepis indica Fauvel, 1928 : 93, fig. 2. g-m; 1953 : 313, fig. 165. g-m. 

Malacoceros indicus: Pettibone, I9&3a : 99 (synonymy); Day, 1967 : 477, fig. 18.5. p-u. 

HABITAT. Silty mud and sand about MTL. 
RECORDS. Nggela Is. - 2 ; Tetel Is. - i ; Fintry Pt. - 9. 

NOTES. Specimens have 10 to 12 bidentate hooks in the neuropodia; in specimens 
from India there are less than 6. 

DISTRIBUTION. Indo-west-Pacific. 



Nerinides sp. cf. gilchristi Day, 1961 

Nerinides gilchristi Day, 1961 : 491, fig. 5. a-d; 1967 : 485, fig. 18.7. i-1. 
HABITAT. Silty sand (Thalassia flat). 
RECORD. Komimbo Bay - i. 

NOTES. The specimen is in two parts - an anterior fragment of 15 setigers and a 
middle body section of 13 setigers. In the latter, the branchiae are flag-shaped, 
corresponding to those of N. gilchristi. However there are 7 or 8 hooded hooks per 
neuropodium, each with a single tooth above the main fang, not 10 to 12 hooks having 
three teeth above the main fang, as in N. gilchristi. 



Polydorella novaegeorgiae n. sp. 

(Fig. 12. A-F) 

DESCRIPTION. All five specimens are incomplete. The largest fragment, chosen 
for the holotype, measures 15 mm for 50 setigers and has a width of i-o to 1-5 mm. 
The colour is whitish except for a brownish pigmentation on the peristomium and 
anterior segments. 

The prostomium has a rounded anterior margin and extends back as a low ridge 
to the middle of setiger 2. On its dorsal surface there are two pairs of eyes and a 
small occipital papilla (fig. 12. A). The broad peristomium carries a pair of stout 
palps which reach to about setiger 15. 

On the anterior segments, the notopodial lobes gradually increase in size and are 
largest on setiger 4. The neuropodial lobes of setigers i to 6 are similar in size and 
are semi-circular in outline; from setiger 7 they are small and inconspicuous. 
Branchiae commence on setiger 6 and continue to setigers 15 to 18 (i.e. there are 10 
to 13 pairs). These are cirriform in shape, the longest being equal to the body width. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 



169 



Both notosetae and neurosetae in the anterior segments, with the exception of the 
notosetae of setiger 4, are winged capillaries. The notopodial bundles are composed 
of two rows of curved capillaries which have narrow blades on one side (fig. 12. c), 
plus a superior group of similar form but much longer. The neurosetae are narrow- 
winged capillaries, each finely tapering to a sharp point. 

As is characteristic of the genus, the notopodium of setiger 4 carries modified 
setae. Two types are present in a double row that follows a wide arc (fig. 12. B) ; 
those of the anterior row are strongly angled at their distal end with a broad blade 
showing fine striations (fig. 12. D) while those of the posterior row are stout hooks 
with bluntly tapering, slightly curved tips (fig. 12. E). There are between 20 and 
25 pairs of the two types of modified setae, plus a superior group of about 15 narrow- 
winged capillaries, in each of the notopodia of setiger 4. In the post-branchial 
region the superior capillary group is replaced by a group of about 10 needle-like 
setae. Bidentate hooded hooks (fig. 12. F) commence at setiger 7 and number 
between 18 and 20 per neuropodium. 

The type species Polydorella prolifera Augener was described from Sharks Bay, 
Western Australia, (Augener, 1914) and has been recorded from India (Gravely, 






FIG. 12. Polydorella novaegeorgiae n.sp. (A) Dorsal view of anterior region. (B) Lateral 
view of setiger 4. (c) Notopodial capillary. (D-E) Modified notopodial setae from 
setiger 4. (F) Neuropodial hook. 



iyo P. E. GIBBS 

1927; Fauvel, 1930). It differs from P. novaegeorgiae in many respects, particularly 
in the form of the prostomium, the structure of the modified setae of setiger 4 and 
in the number of branchiae. A further point of difference is that the specimens 
from Marovo Lagoon do not show any sign of reproduction by scissiparity, a feature 
observed in P. prolifera. 

HABITAT. Mud at n m depth. 

RECORDS. ML 39-5. 

British Museum (Natural History) Registration No. Holotype 1970-55 

Paratypes 1970-56 



Prionospio cirri/era Wire"n, 1883 

Prionospio cirrifera : Fauvel, 1927 : 62, fig. 2i.k-n; 1953 : 324, fig. 164. k-m; Day, 1967 : 486, 
fig. 18.8. a-d. 

HABITAT. Mud at 22 m depth. 

RECORD. ML 29-1. 

NOTES. Although P. cirrifera has not been recorded previously from the west 
Pacific region the single specimen agrees in detail. It has 12 pairs of branchiae and 
the hooded crotchets are multidentate, corresponding to the northern form (cf. Day, 
1967). 

DISTRIBUTION. North Atlantic; North Pacific; India; South Africa. 



Prionospio ehlersi Fauvel, 1928 
Prionospio ehlersi Fauvel, ig28a :'io, fig. i. a-r; Day, 1967 : 490, fig. 18.9. d-f. 

HABITAT. Mud and sand, 2-22 m depth. 

RECORDS. ML 37 - 2 ; ML 41 - 4 ; ML 134 - 1 ; ML 156- 8 ; ML 191 - 1 ; ML 194 -2 ; 
ML 195-1; ML 218 -6. 

NOTE. This is the first record of P. ehlersi from the Pacific region. 

DISTRIBUTION. Morocco; East Africa. 



Prionospio malmgreni Claparede, 1870 

Prionospio malmgreni: Fauvel, 1927 : 61, fig. 21. a-e; Hartman & Imajima, 1964 : 285; Day, 
1967 : 492, fig. 18.9. a-c. 

HABITAT. Silty sand at 2 m depth. 
RECORDS. ML 196 - 2. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 171 

NOTES. Both specimens are small, about 10 mm long. Across setiger 7 there is 
a membraneous ridge and this feature separates P. malmgreni from the closely allied 
species P. steenstrupi Malmgren. 

DISTRIBUTION. Atlantic; Mediterranean; South Africa; South-west Australia; 
Japan; southern California. 



Prionospio pinnata Ehlers, 1901 

Prionospio pinnata : Fauvel, 1953 : 323, fig. 174. e; Imajima & Hartman, 1964 : 286; Day, 1967 : 
488, fig. 18.8. i-1. 

HABITAT. Mud and silty sand, 2-26 m depth 

RECORDS. ML 37-10; ML 40-5; ML 41 -27;. ML no -4; ML 156- 2; ML 
157 - 2 ; ML 191 - i ; ML 194 - 2 ; ML 195 - i. 

DISTRIBUTION. Cosmopolitan. 



Prionospio steenstrupi malayensis Caullery, 1914 

Prionospio steenstrupi malayensis: Caullery, 1944 : 14, fig. 8. a-f. 

HABITAT. A wide variety of deposits from mud to coarse sand, 2-24 m depth. 

RECORDS. ML 55 - 2 ; ML 68 - i ; ML 188 - i ; ML 192 - i ; ML 196 - 5 ; ML 204 - 
i; ML 230- i. 

NOTES. Caullery recognized the subspecies malayensis on the grounds of geo- 
graphical separation from the stem species which was then (1914) known only from 
the North Atlantic. In the Solomon Islands specimens there are low membraneous 
ridges across the dorsum from setiger 12 to setiger 16 or 17. These ridges are not 
present in the stem species and thus this feature distinguishes the two forms. 

DISTRIBUTION. East Indies. 



Prionospio tetelensis n. sp. 

(Fig. 13. A-D) 

DESCRIPTION. All specimens are incomplete. The holotype from ML 188 is 
14 mm long for 42 segments and has a diameter of 2 to 3 mm. One of the paratypes 
(from ML 68) is a mature female containing oocytes which are turquoise blue in 
colour. 

The prostomium is rounded anteriorly and extends back to setiger 3 as an elevated 
keel between peristomial folds (fig. 13. B). There are three pairs of prostomial 



I 7 2 



P. E. GIBBS 



eyes, two pairs of which are small and situated to the anterior of a larger, transverse 
pair. On either side of the prostomium, the peristomium is extended to form lateral 
wing-like expansions. 

The first setiger is small and lacks notosetae. The parapodial lobes of the anterior 
segments are large and ear-shaped, the pre-setal ones being smaller than the post- 
setal (fig. 13. c). These lobes gradually decrease in size, becoming flatter and less 
rounded in the middle body. From setiger 6 to between setigers 20 to 26 the post- 
setal notopodial lobes are joined across the dorsum by membraneous ridges. In 
mature specimens there are no genital pockets between the anterior neuropodia. 

Branchiae are present on setigers 2 to 5 and all are pinnate (fig. 13. A). Each has 
a short stem and numerous pinnules arising from the lateral and posterior surfaces 
of the main axis. The size of the branchiae decreases from the first to the fourth 
pair. 

Both notosetae and neurosetae in the anterior segments are narrow-winged capil- 
laries. Commencing between setigers 20 and 22, about 12 hooks appear in the 
neuropodia, accompanied by very fine pointed setae and also 2 or 3 sabre-like setae. 
Hooks do not appear in the notopodia until about setiger 50. Each hook is hooded 
with a series of five teeth above the main fang (fig. 13. D). 






FIG. 13. Prionospio tetelensis n.sp. (A-B) Lateral and dorsal views of anterior 
region, (c) Parapodium of setiger 15. (D) Neuropodial hook. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 173 

Of the many species of the genus Prionospio now known, three are similar to P. 
tetelensis in possessing four pairs of pinnate branchiae on setigers 2 to 5. These are 
P. treadwelli Hartman, P. tennis (Verrill) - both from the Atlantic coast of North 
America - and P. peruana Hartmann-Schroder from Peru. P. tetelensis may be 
distinguished from all three species on the basis of a combination of characters, 
namely, the prostomial keel between peristomial folds extending to setiger 3, the 
lateral wing-like expansions of the peristomium, the membraneous ridges across the 
dorsum of setigers 6 to 20 and the neuropodial hooks being present from setiger 20. 

HABITAT. Mud from LWM to 18 m depth. 

RECORDS. Tetel Is. - i ; ML 68 - i ; ML 188 - i (holotype) ; ML 195 - i ; ML 
228 - 1. 

British Museum (Natural History) Registration No. Holotype 1970-57 

Paratypes 1970-58-61 



Pseudopolydora corallicola Woodwick, 1964 

Pseudopolydora corallicola Woodwick, 1964 : 151, fig. 2. 9-12. 

HABITAT. Coralline mud at LWM ; silty sand at 2 m depth. 
RECORDS. Matiu Is. - i ; ML 196 - i. 

NOTES. Both specimens are small (under 5 mm long) and are damaged. Those 
characters that can be checked, such as the prostomial ridge extending to setiger 6, 
the modified setae of setiger 5 and the peculiar fenestrated neuropodial hooks, all 
correspond. 

DISTRIBUTION. Marshall Islands. 



Pseudopolydora sp. 

HABITAT. In brackish water (5-6 % ) - boring in sponge. 

RECORDS. Lagoon at the mouth of the Lunga River at Lunga Point - numerous 
specimens. 

NOTES. These specimens are up to 5 mm in length. The modified setae of 
setiger 5 include an anterior row of broad-winged, sharply tapering setae and a 
posterior row of stout hooks with recurved tips: there are 6 or 7 pairs of the two 
types. Branchiae commence on setiger 6 and extend to setigers 16 or 17 (n or 12 
pairs). Neuropodial hooks are present from setiger 8. In the posterior notosetae 
there are one or two stout, curved hooks which arch over the dorsum. 

Mr. W. J. Light has examined this series and, in a personal communication, he 



174 P. E. GIBBS 



has pointed out that certain characters, in particular the form of the modified setae 
on setiger 5 and also the presence of specialized posterior notosetae, indicate that 
these specimens represent a new species of Pseudopolydora. He has kindly consented 
to publish a description in a separate paper. 



Scolelepis squamata mendanai n. subsp. 

DESCRIPTION. The holotype is a complete specimen measuring 50 mm in length 
for about 200 segments. The diameter of the body is between i-o to 1-5 mm. The 
palps are very small and slender extending only to setiger 3 or 4. There are four 
eyes set in a transverse row across the prostomium. Branchiae commence on setiger 
2 and are fused to the notopodial lamellae for about one-third of their length. 
Bidentate hooks number 9 or 10 in the neuropodia from about setiger 45, and I to 3 
in the notopodia starting between setigers 90 to 100. The post-setal neuropodial 
lobes are ear-shaped in anterior segments but become bilobed between setigers 35 to 
45 and are distinctly divided in the middle segments. 

The subspecies mendanai is distinct from the stem species (Fauvel, 1927 : 36, 
fig. ii. g-n, as Nerine cirratulus) in the following characters: (i) the palps are very 
small reaching only to setiger 3 (not setiger 24), (ii) neuropodial hooks are present 
from setiger 45 (not setiger 40) and (iii) notopodial hooks are present from about 
setiger 90 (not setiger 60). The subspecies saipanensis (Hartman, 19540 : 230, fig. 
2. a-e, as Nerine cirratulus saipanensis) differs in that the hooks appear in the neuro- 
podia from setigers 29 to 31 and in the notopodia from setiger 41. 

HABITAT. Silty sand, MTL to LWM. 

RECORDS. Graham Pt. - 5 ; Fintry Pt. - 6 (including holotype). 
British Museum (Natural History) Registration No. Holotype 1970-62 

Paratypes 1970-63-65 



Spio filicornis (Muller, 1776) 

Spiofilicornis: Fauvel, 1927 : 43, fig. 15. a-g; Imajima & Hartman, 1964 : 289; Day, 1967 : 481, 
fig. 18.6. l-o. 

HABITAT. Coralline mud at LWM. 

RECORDS. Matiu Is. - 8. 

DISTRIBUTION. North Atlantic ; South Africa ; North Pacific. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 175 

Family MAGELONIDAE 
M agelona japonica Okuda, 1937 

Magelona japonica Okuda, 1937 : 247, figs 23-24; Imajima & Hartman, 1964 : 290; Tampi & 
Rangarajan, 1964 : 113, figs 44-49. 

HABITAT. Coarse sand below LWM. 

RECORDS. Tetel Is. - 2. 

DISTRIBUTION. Bay of Bengal; North Pacific. 



Magelona sp. 

HABITAT. Silty sand at 2 m depth. 
RECORD. ML 283 - i. 

NOTES. The specimen is an anterior fragment of 20 segments. The prostomium 
is elongated with a rounded anterior margin lacking frontal horns. Anterior segments 
have large notopodial and small neuropodial lamellae. The setae of segment 9 are 
unspecialized. From setiger 10 the parapodial lamellae are rhomboidal. The 
abdominal hooded hooks are bidentate with a single tooth above the main fang. 

The specimen is close to M. capensis Day, differing in the shape of the abdominal 
lamellae being rhomboidal not oval and in the hooks being bidentate not tridentate. 



Family TROCHOCHAETIDAE 
Poecilochaetus serpens honiarae n. subsp. 

DESCRIPTION. All of the specimens are incomplete; the largest specimen (the 
holotype) is 20 mm long with 44 segments and has a body width of 2-0 mm. The 
characters are those of the stem species (Allen, 1904 : 79, pi. 7-12, text-fig, i) except 
that the aristate spines, with knobbed ends and hairy terminal aristae (Allen, 1904 : 
pi. 9, fig. 18) commence between setigers 20 to 25 and thus are not confined to the 
posterior region beyond setiger 80, as in the stem species. 

HABITAT. Silty and coarse sand, MTL to LWM. 

RECORDS. Tetel Is. - 3 (including holotype) ; Komimbo Bay - 2. 
British Museum (Natural History) Registration No. Holotype 1970-66 

Paratypes 1970-67-69 

NOTES. The Solomon Islands specimens have a reddish, chitinous, triangular 
plate on the dorsum of setiger 9 (a similar, but diamond-shaped, structure has been 



176 P. E. GIBBS 

noted in P. johnsoni by Hartman, 1939). Lateral sense organs are missing from 
setigers 6 to 9 and the branchiae, composed of two filaments on the posterior sides 
of the parapodia, are present from setigers 17 or 18. 

An examination of P. serpens individuals from the type locality at Plymouth 
reveals that Allen overlooked the dorsal plate on setiger 9 and also the absence of 
lateral sense organs on setigers 6 to 9. Allen states that there are two pairs of 
branchial filaments from setiger 21 but in the several specimens which have been 
examined there is only one pair, although a third filament sometimes occurs after 
setiger 32. 

Thus adult specimens of the stem species P. serpens from the type locality and 
adults of the subspecies honiarae from the Solomon Islands are very similar, the 
distinguishing feature being the presence of aristate spines in the middle body seg- 
ments of the latter. This distinction is recognized as a minor one, deserving of sub- 
specific separation only. 

P. serpens is remarkable in its retention of the pelagic habit until a late stage of 
larval development. Thus it is not surprising that the existence of this species in the 
Indo-west Pacific region was known from plankton records long before adult speci- 
mens were discovered in the region. Pelagic stages have been described from Indo- 
China (Fauvel, 1939) and India (Banse, 1959; Ganapati & Radhakrishna, 1958). 
Only recently has an adult specimen been discovered in Indian waters (Achari, 1968). 



Poecilochaetus tropicus Okuda, 1935 

Poecilochaetus tropicus Okuda, 1935 : 289, figs 1-2; I937a : 294, figs 39-40. 
HABITAT. Silty and coarse sand from MTL to LWM. 
RECORDS. Tetel Is. - 5; Graham Pt. - i. 

NOTES. P. tropicus was described from an incomplete specimen of 39 segments 
taken in the Palau Islands in 1934. No further material appears to have been 
recorded since that date. 

All of the Solomon Islands specimens are incomplete. The most complete 
individual is immature, measuring 26 mm long for 70 setigers, but the largest specimen 
appears mature and has a length of 30 mm for 50 setigers. The original description 
can be supplemented with the following details - (i) the two branchial filaments 
situated on the posterior sides of the parapodia are not present in small specimens 
but are conspicuous in mature individuals; (ii) on the dorsal surface of setiger 9, 
there is a small, yellowish, semi-lunar-shaped plate (a similar structure to that in 
P. serpens)', (iii) the flask-shaped parapoch'al lobes of setigers 7 to 13 contain an 
iridescent, light blue material in mature specimens, (iv) the spines with hairy 
terminal filaments (Okuda, 1935, fig. 2.e) are not replaced by the knobbed, aristate 
type (as in P. serpens) in the region anterior to setiger 70 in the most complete 
specimen available. 

DISTRIBUTION. Palau Islands. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 177 

Family CHAETOPTERIDAE 
Chaetopterus variopedatus (Renier, 1804) 

Chaetopterusvariopedatus: Fauvel, 1927 : 77, fig. 26. a-n; 1953 : 337, fig. 175. a-n; Day, 1967 : 
529, fig. 22.2. a-g. 

HABITAT. Attached to Acropora on wharf piles; in sand at 2 m depth. 
RECORDS. Yandina - 5 ; ML 118-2. 

NOTE. All of the specimens are small, the largest being only 20 mm long. 
DISTRIBUTION. Cosmopolitan. 



Mesochaetopterus Sagittarius (Claparede, 1870) 

Mesochaetopterus minutus Potts, 1914 : 963, pi. 2, fig. 4, pi. 3, figs 7-8, text-figs 4-5; Fauvel, 

1953 : 342, fig- i? 8 - a; Da Y. J 96? : 53L fi g- 22.2. h-n. 
Mesochaetopterus Sagittarius: Bhaud, 1969 : 325, fig. i. a-f. 

HABITAT. A wide variety of deposits from coralline mud to coarse sand, MTL to 
LWM ; in Halichondria at 5 m depth. 

RECORDS. Tetel Is. - 25 ; Komimbo Bay - 22 ; Matiu Is. - i ; Maraunibina Is. - 2 ; 
Graham Pt. - i ; Pigeon Is. - num. ; Fintry Pt. - 8 ; Yandina - num. 

NOTES. The type specimens measure 25 mm long with a diameter of i mm 
(Potts, 1914) but Monro (1931) records specimens from N.E. Australia that are twice 
this size (50 x 2 mm) with tubes up to 20 cm in length. Some of the specimens 
from Tetel Is. and Komimbo Bay, and all of those from Pigeon Is. and Yandina are 
similar in size to the type material but the remaining specimens are larger and 
approximate the size of the Barrier Reef specimens. 

DISTRIBUTION. Mediterranean ; Indo-west-Pacific to Japan. 



Phyllochaetopterus elioti Crossland, 1903 

Phyllochaetopterus elioti Crossland, 1903 : 172, pi. 16, figs 1-4, 7-8, pi. 17, figs 10-13; Fauvel, 
1953 : 34. fi g- 177- e - h : Da Y. 1967 : 525. fi g- 22.1. f-g. 

HABITAT. Abundant in silty sand, MTL to LWM ; coarse sand at 4 m depth. 
RECORDS. Tetel Is. - num. ; Fintry Pt. - num. ; ML 295 - i. 
DISTRIBUTION. South and East Africa; India. 



178 P. E. GIBBS 

Phyllochaetopterus herdmani (Hornell, 1903) 

Phyllochaetopterus herdmani: Willey, 1905; 292, pi. 5, figs 127-132; Fauvel, 1953 : 342, fig. 177. 
i-m; Day, 1967 : 524, fig. 22.1. a-e. 

HABITAT. Silty sand at MTL. 
RECORD. Tetel Is. - i. 

NOTE. The single specimen is rather damaged and in a poor state of preservation 
but nevertheless corresponds to the species description. It is anterior fragment of 
31 setigers comprising n anterior, 2 middle and 18 abdominal segments. On setiger 
4 there are 9 modified setae in each of the notopodia. This is a first record for the 
Pacific region. 

DISTRIBUTION. East Africa; Ceylon. 



Phyllochaetopterus socialis Claparede, 1870 

Phyllochaetopterus socialis : Fauvel, 1927 : 84, fig. 30. a-1; 1953 : 339, fig. 176. a-1; Day, 1967 : 
525, fig. 22.1. h-r. 

HABITAT. Very abundant on moderately exposed reef platforms, forming dense 
colonies towards LWM (see Gibbs, 1969, fig. 136); single specimens embedded in 
Milkpora and the sponge Neofolitispa dianchora (de Laubenfels) from 5 m depth. 

RECORDS. Matiu Is., Cape Esperance and Batuona Is. - numerous specimens; 
Maramasike Pg. - 3; Kokomtambu Is. - i ; Yandina - i. 

DISTRIBUTION. Mediterranean; Atlantic; Indo-Pacific. 



Spiochaetopterus costarum costarum (Claparede, 1868) 

Telepsavus vitrarius Ehlers, 1908 : 114, pi. 15, figs 1-8. 
Spiochaetopterus vitrarius: Day, 1967 : 528, fig. 22.1. u-v. 
Spiochaetopterus costarum costarum: Gitay, 1969 : 14 (synonymy). 

HABITAT. Silty sand (Thalassia flat) at about MTL ; coarse sand at LWM. 
RECORDS. Tetel Is. - 6 ; Komimbo Bay - 14. 

NOTES. The specimens have a rounded prostomium with a pair of lateral eyes. 
The anterior region is composed of 9 setigers, the notopodia of setiger 4 bearing single 
modified setae with swollen, triangular or heart-shaped tips. On the ventral 
surface of setigers 7 to 9 the tissue is distinctly glandular, being brownish-black in 
colour on setiger 7 but whitish on setigers 8 and 9. On setiger 2 there is usually a 
thin line of brownish pigment across the ventrum. Up to 19 middle and 25 ab- 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 179 

dominal segments have been noted in several specimens but most of the material is 
poorly preserved. The largest specimens measure about 25 mm in length, their 
tubes being up to 60 mm long and 0-5 to i-o mm in diameter. The latter are trans- 
parent and highly wrinkled. 

Gitay (1969) has recognized three subspecies of 5. costarum in the Pacific region, 
namely pottsi, monroi and okudai. The specimens from the Solomon Islands differ 
from all three in having fewer middle segments (i.e. 19 compared to 30-90, about 50 
and about 67 respectively). Furthermore their small size distinguishes them from 
pottsi and the presence of eyes separates them from monroi and okudai. However 
these specimens appear to be identical with T. vitrarius Ehlers, known from the 
eastern Atlantic and Natal, which Gitay considers to be a synonym of S. costarum 
costarum. 

DISTRIBUTION. Eastern Atlantic; Mediterranean; Indian Ocean. 



Family GIRRATULIDAE 
Cirriformia filigera (Delle Chiaje, 1825) 

Audouinia filigera: Fauvel, 1927 : 92, fig. 32. h-m; 1953 : 331, fig. 173. h-1. 
Cirriformia filigera Day, 1967 : 518, fig. 20.4. p-q. 

HABITAT. In Acropora rubble and under coral boulders on reef platforms; in 
crevices and vacated borings in Porites, Acropora and beachrock. 

RECORDS. Kokomtambu Is. - i ; Tetel Is. - 3 ; Komimbo Bay - 3 ; Matiu Is. - i ; 
Paleki Is. - 2; Lauvie Is. - 2. 
DISTRIBUTION. Atlantic; Mediterranean; Indian Ocean. 



Cirriformia punctata (Grube, 1859) 

Audouinia semicincta: Fauvel, 1953 : 330 fig. 174. c; Okuda, 1937 : 2 96, fig. 41. a-c. 
Cirriformia punctata: Day, 1967: 517, fig. 20.4. j-m. 

HABITAT. In sedimented crevices on reef platform. 
RECORDS. Komimbo Bay - i ; Batuona Is. - 54. 
DISTRIBUTION. Circumtropical. 



Dodecaceria fistulicola Ehlers, 1901 

Dodecaceria fistulicola : Fauvel, 1953 : 335, figs 169. h-i, 174. a-b; Okuda, 1937 : 2 98, fig. 42. a-f. 
HABITAT. Crevices in reef platform. 
RECORDS. Komimbo Bay - i ; Matiu Is. - i. 



180 P. E. GIBBS 

NOTES. A complete specimen has 68 segments and is about 15 mm long. There 
are 4 or 5 pairs of branchiae which are shorter than the palps. The first 7 or 8 
setigers carry capillary setae which are replaced in later segments by 5 or 6 hooks in 
both rami. 

DISTRIBUTION. Indo-west-Pacific ; southern South America. 



Dodecaceria laddi Hartman, 1954 

Dodecaceria laddi Hartman, 1954 : 638, n 8 s I 7^- c > J 77- d-h; Day, 1967 : 502, fig. 20.1. g-i. 

HABITAT. Boring (?) in Forties. 
RECORD. Kokomtambu Is. - i. 
DISTRIBUTION. Marshall Islands; South Africa. 



Tharyx sp. 

HABITAT. Silty sand at 36 m depth. 
RECORD. Off Fintry Pt. - i. 

NOTES. The specimen is incomplete with 26 segments and measures 3-5 mm long. 
The diameter is about 0-4 mm. The palps have been lost but scars indicate their 
position. The setae are entirely capillary. It is impossible to identify this fragment 
with any certainty but possibly it is T. multifilis Moore, which is known from Madras 
(Fauvel, 1953). 



Family GOSSURIDAE 
Cossura coasta Kitamori, 1960 

Cossura coasta Kitamori, 1960 : 1082, fig. i. a-f; Imajima & Hartman, 1964 : 299; Day, 1967 : 
581, fig. 26.1. a-d. 

HABITAT. Mud at 22 m depth. 

RECORD. ML 37 - 2. 

NOTES. Both specimens are incomplete; the larger measures 10 mm for 60 
segments and has a diameter of 0-5 mm. They differ from the species description 
only in respect of the shorter capillary setae which, instead of being serrated, are 
smooth, and also taper quite abruptly about halfway along their length to a very 
slender filament. 

DISTRIBUTION. Southern Africa; Japan. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 181 

Family FLABELLIGERIDAE 
Diplocirrus glaucus orientalis n. subsp. 

DESCRIPTION. The specimen chosen for the holotype (from ML 194) is 18 mm long 
for 25 segments and has a maximum diameter of about 2-0 mm in the anterior body 
region. A complete paratype (from ML 55) is 20 mm in length for 40 segments. 
The characters are those of the stem species Diplocirrus glaucus (Malmgren), (Fauvel, 
1927 : 120, fig. 43. a-d) but is distinguished by the presence of conspicuous orange 
coloured, globular papillae situated on the ventral surface below each neuropodium 
from setiger 4 to setigers 14 or 16. 

HABITAT. Mud and silty sand, n to 24 m depth. 

RECORDS. ML 55 - i ; ML 188 - 3 ; ML 194 - i (holotype) ; ML 218 - i ; 
ML 230 - 1. 

British Museum (Natural History) Registration No. Holotype 1970-70 

Paratypes 1970-71-74 

NOTES. The stem species is widely distributed in the North Atlantic region and, 
although Fauvel (1932) refers a specimen dredged in the Mergui Archipelago from 
7 m (4 fm) to this species 'with great hesitation', no certain record of this species 
being taken in the Indian Ocean exists. 

Specimens of the stem species from Plymouth do not possess conspicuous papillae 
below the neuropodia of the anterior segments. The presence of these papillae thus 
distinguishes the subspecies orientalis. 



Family SGALIBREGMIDAE 
Hyboscolex longiseta Schmarda, 1861 

Hyboscolex longiseta Schmarda, 1861 : 54, pi. 27, fig. 211, with text-figs; Day, 1967 : 588, fig. 
27.2. a-d. 

HABITAT. Under coral boulders in muddy silt ; in beachrock. 
RECORDS. Tetel Is. - i ; Maraunibina Is. - i ; Lauvie Is. - 2. 
DISTRIBUTION. Southern and East Africa; New Zealand (Auckland Is.). 



Scalibregma inflatum Rathke, 1843 

Scalibregma inflatum: Fauvel, 1927 : 123, fig. 44. a-f; 1953 : 355, fig. 185. a-f; Day, 1967 : 590, 
fig. 27.2. e-i. 

HABITAT. Silty mud at 13 m depth. 

RECORD. ML 69-1. 

DISTRIBUTION. Cosmopolitan (but few records from the Indo-Pacific region). 



i8a P. E. GIBBS 

Family OPHELIIDAE 
Armandia lanceolata Willey, 1905 

Armandia lanceolata Willey, 1905 : 288, pi. 5, fig. 120; Fauvel, 1953 : 358. 

HABITAT. Acropora rubble at LWM; silt and sand, 5-24 m depth. 

RECORDS. Tetel Is. -i; ML 56-1; ML 117 - i; ML 134 - i; ML 229 - i; 
ML 230 - 1. 

DISTRIBUTION. Indo-west-Pacific. 



Armandia leptocirrus (Grube, 1878) 

Ophelina (Armandia) leptocirrus Grube, 1878 : 194. 

Armandia leptocirrus: Fauvel, 1953 : 358; Day, 1967 : 577, fig. 25.2. h. 

HABITAT. Silt and coralline mud at LWM ; mud and sand, 2-18 m depth. 

RECORDS. Komimbo Bay - i ; Matiu Is. - 7 ; ML 68 - 2 ; ML 97 - i ; ML 118 - i 
ML 190 - 2 ; ML 192 - i ; ML 196 - 3 ; ML 283 - 4. 

DISTRIBUTION. Tropical Indo-west-Pacific. 



Armandia longicaudata (Caullery, 1944) 

Ammotrypane longicaudata Caullery, 1944 : 44, fig. 35. a-c. 
Armandia longicaudata: Day, 1967 : 577, fig. 25.2. a-c. 

HABITAT. Silty sand at 24 m depth. 

RECORD. ML 230 - i. 

DISTRIBUTION. East Africa; East Indies. 



Ophelia koloana n. sp. 

(Fig. 14. A-B) 

DESCRIPTION. The two type specimens measure 10 to 12 mm in length and have 
a diameter between 1-0-1-5 mm. The paratype is a female approaching maturity, 
containing coelomic oocytes. 

The prostomium is sharply pointed. The anterior region is cylindrical with the 
ventral groove commencing at setiger 8. The first 9 setigers are abranchiate, 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 183 

branchiae being present from setiger 10 to setiger 27 (i.e. there are 18 pairs). Each 
branchia is rather small and has a 'crimped' appearance (fig. 14. A). The branchi- 
ferous region is followed by 4 abranchiate segments and then two achaetous annuli 
(or pre-anal segments) . Thus, following the scheme of Tebble (1953) , the body formula 
is ga + i8b ( ) + 4a + 2n = 33. From setiger 30 to the pygidium there is a pair 
of prominent dorso-lateral ridges (fig. 14. B). The pygidium bears two stout ventral 
papillae and 9 small dorsal ones. Neither nephridiopores nor branchial fenestrations 
in the lateral body wall could be detected. 

In terms of its affinities, 0. koloana has a similar body formula to 0. denticulata 
Verrill and 0. capensis Kirkegaard. However in 0. denticulata, which is an Atlantic 
species, the ventral groove starts at segment 10 and it has n to 18 dorsal anal 
papillae (Fauvel, 1927 : 132, fig. 46, g-h, as 0. neglecta, fide Tebble, 1953). 0. 
capensis from South Africa differs in that it has lateral swellings from segment 26 to 
the pygidium (Kirkegaard, 1959 : 45, fig. 8) not dorso-lateral ridges from setiger 29. 

HABITAT. Sand at 2 m depth. 

RECORDS. ML 118-2. 

British Museum (Natural History) Registration No. Holotype 1970-75 

Paratype 1970-76 

NOTE. Hitherto the genus Ophelia appears to have been recorded only once in 
the tropical Indo-west-Pacific region, namely, a single specimen of 0. limacina from 
Indo-China (Fauvel, 1939). 





0-5mm 0-5mm 

FIG. 14. Ophelia koloana n.sp. (A) Lateral view of segments 22 to 24. 
(B) Dorso-lateral view of posterior region. 



Polyophthalmus pictus (Dujardin, 1839) 

Polyopht halmus pictus : Fauvel, 1927 : 137, fig. 48. l-o; 1953 : 3^> & 1 ^7- l-o; Day, 1967 : 579, 
fig. 25.2. k-m. 

HABITAT. Interstices of the sponge Halichondria at 5 m depth. 
RECORDS. Yandina - 3. 



184 P. E. GIBBS 

NOTES. All three specimens appear to be juvenile in that they measure only 
3 to 7 mm in length. Pillai (1965) found this species in a similar habitat, namely, 
within sponges and other encrusting organisms on bamboo oyster spat collectors. 

DISTRIBUTION. Cosmopolitan. 



Family STERNASPIDAE 
Sternaspis scutata (Renier, 1807) 

Sternaspis scutata: Fauvel, 1927 : 216, fig. 76. a-g; 1953 : 401, fig. 210. a-g; Day, 1967 : 648, 
fig. 31.1. a-d. 

HABITAT. Mud and silty sand, 18-24 m depth. 
RECORDS. ML 218 - 6 ; ML 230 - 3. 
DISTRIBUTION. Cosmopolitan. 



Family CAPITELLIDAE 
Capitellethus dispar (Ehlers, 1907) 

Capitellethus dispar: Fauvel, 1953 : 37*^ Rullier, 1965 : 194. 

HABITAT. Silty sand above LWM. 
RECORDS. Komimbo Bay - 3. 

NOTES. The material is fragmentary and the largest specimen measures 18 mm 
long for 60 segments. The'peristomium is achaetous and there are n thoracic 
setigers with capillary setae in both rami, which are replaced by hooks from setiger 
12. No branchiae could be seen. In all three specimens the epidermal cells of 
setigers n and 12 (the last thoracic and first abdominal) contain a dark pigment. 

DISTRIBUTION. India; West Pacific. 



Capitobranchus sp. 

HABITAT. Coarse sand at LWM. 
RECORD. Tetel Is. - i. 

NOTES. The single specimen is incomplete, measuring 20 mm for 20 segments, 
and has a diameter of about 2 mm. The peristomium is biannulate, segments 2 to 
4 are triannulate, segments 5 to 8 are quadriannulate and the remaining segments are 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 185 

biannulate. Segments 2 and 3 are achaetous but may have lost their setae. Seg- 
ments 4 to 12 have capillaries in both rami which are replaced by hooks in the 
neuropodia from segment 13. 

In having at least 19 thoracic segments and hooks from setiger 13 (assuming 
segments 2 and 3 to have been setigerous), the specimen falls within the broad 
definition of the genus Capitobranchus given by Day (1962, p. 651) as follows: 
"Thorax with an achaetous peristome followed by about 19 setigerous segments 
most of which bear capillary setae in both rami, but hooded hooks are present in the 
last few neuropodia." The genus is monospecinc based on C. macgregori from 
Madagascar which has 19 thoracic setigers, the last four of which bear neuropodial 
hooks. Due to its incomplete nature, the specimen from Tetel Island must remain 
indeterminable as to species until further material becomes available. 

Dasybranchus caducus (Grube, 1846) 

Dasybranchus caducus: Fauvel, 1927 : 148, fig. 52. a-h; 1953, 365, fig. 190. a-h; Day, 1967 : 603, 
fig. 28.3. e-h. 

HABITAT. A wide variety of deposits, mud to coarse sand, particularly under coral 
boulders on reef platforms, from above MTL to LWM ; mud, 9-22 m depth. 

RECORDS. Tetel Is. - 56; Nggela Is. (Sandfly Pg.) - 30; Naro Bay - 7; Komimbo 
Bay - 14; Matiu Is. - 16; Graham Pt. - 19; Fintry Pt. - 7; ML 29-2; ML 56 - I. 

DISTRIBUTION. Atlantic; Mediterranean; Indo-Pacific. 



Mastobranchus dollfusi Fauvel, 1936 
Mastobranchus dollfusi Fauvel, 1936 : 81, fig. n. a-g; Kirkegaard, 1959 : 51. 

HABITAT. Muddy sand at MTL; mud and silty sand, 2-22 m depth. 

RECORDS. Nggela Is. - 4 ; ML 29 - i ; ML 68 - 5 ; ML 195 - i. 

NOTES. Specimens have a diameter of 0-5 to i-o mm and the largest anterior 
fragment is 30 mm long for 70 segments. The prostomium is conical with a pair of 
pigmented eye-spots and is followed by an achaetous peristomium. The first setiger 
has capillary notosetae only; setigers 2 to 10 have capillaries in both rami. The 
number of thoracic setigers varies, as noted by both Fauvel and Kirkegaard for the 
West African material. In the Solomon Islands specimens setiger n usually has 
capillary notosetae and neuropodial hooks with the succeeding segments carrying 
hooks in both rami. However two specimens have capillary notosetae on setiger 12, 
one of which also has capillary neurosetae on setiger n. A fragment of the posterior 
region possesses retractile branchiae situated posteriorly to the notopodia of the 
abdominal segments, and the pygidium has four short anal cirri. 

Despite the wide geographical separation of the two localities, the Solomon Islands 
specimens appear to be identical to those from West Africa in all details. 

DISTRIBUTION. West Africa. 



186 P. E. GIBBS 

Mastobranchus trinchesii Eisig, 1887 

Mastobranchus trinchesii: Fauvel, 1927 : 152, fig. 54. a-i; Kirkegaard, 1959 : 52. 

HABITAT. Silty sand, MTL to LWM. 

RECORDS. Tetel Is. - i ; Komimbo Bay - 4; Graham Pt. - 27. 

NOTES. Although M. trinchesii is a species hitherto known only from the 
Mediterranean and West Africa, the Solomon Islands specimens appear to be identical. 
Briefly, the latter specimens have an achaetous peristomium, followed by n thoracic 
setigers with capillary setae in both rami, except the first which has notosetae only. 
From setiger 12 the capillaries are replaced by hooks in the neuropodia, but in the 
notopodia capillaries persist with the hooks until about setiger 90. Retractile 
cirriform branchiae are present in the posterior segments and the pygidium carries 
four anal cirri. 

DISTRIBUTION. Mediterranean; West Africa. 



Mediomastus sp. cf. capensis Day, 1961 

Mediomastus capensis Day, 1961 : 518, fig. u. a-d; 1967 : 600, fig. 28.2. n-p. 

HABITAT. Mud at 18 m depth. 

RECORD. ML 218 - i. 

NOTES. The specimen is an anterior fragment of 25 segments, measuring 7 mm 
in length. An achaetous peristomium is followed by 10 thoracic setigers, the first 
four of which have capillaries, the remaining six having hooks only. Although 
conforming to the description of the South African material, the identity of the 
specimen must remain provisional, due to its fragmentary condition. 



Notomastus sp. 

HABITAT. Sand, 2-9 m depth. 

RECORDS. ML 97 - i ; ML 118 - i. 

NOTES. Both specimens are anterior fragments consisting of n thoracic setigers 
and a few abdominal segments. Setigers i to 4 are triannulate and setigers 5 to n 
are quadriannulate. The thoracic segments are strongly areolated, particularly 
the more anterior ones. The abdominal tori are well developed. A specific identi- 
fication cannot be made with any certainty because of the absence of branchiferous 
segments. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 187 

Family ARENICOLIDAE 
Abarenicola claparedii claparedii (Levinsen, 1883) 

Arenicola claparedii: Fauvel, 1927 : 163, fig. 57. k-n. 
Abarenicola claparedii: Wells, 1959 : 307, fig. 2. 

HABITAT. Coralline silty mud at LWM. 
RECORD. Matiu Is. - i. 

NOTES. The single specimen was kindly examined by Prof. G. P. Wells who 
identified it, from its superficial characters only, as one of the cystless species of 
Abarenicola, probably claparedii claparedii. This determination will have to be 
confirmed on the basis of its internal anatomy. 

DISTRIBUTION. Mediterranean. 



Family MALDANIDAE 

The following descriptions of the maldanid material are based on notes provided 
by Dr. Charlotte Mangum. 



Subfamily EUCLYMENINAE 
Clymenella (=Macroclymene) sp. 1 

HABITAT. Silty shell gravel on Thalassia flat. 
RECORD. Graham Pt. - i. 

NOTES. The specimen is incomplete (the head and probably five anterior setigers 
are missing) and has 48 setigers plus a caudal funnel. The latter carries long and 
short cirri, which are arranged in an alternating pattern of one long, two short, one 
long, two short and so on, in addition to a median ventral cirrus that is 1-5 to 2-0 
times the length of the long cirri. Posterior pre-anal achaetous segments are absent. 



Clymenella (=Macroclymene) sp. 2 

HABITAT. Under boulder in muddy silt at LWM. 
RECORD. Komimbo Bay- i. 

NOTES. An anterior fragment composed of 27 setigers is available. It has a 
cephalic plaque with nuchal organs extending nine-tenths of its length. Prostomial 
pigment spots are absent. There are shallow clefts in the medial and posterior 
margins of the cephalic rim. Uncini are present on all setigers including the first 
three. 



i88 P. E. GIBBS 

Clymenella (=Euclymene) sp. 

HABITAT. Silty sand with coral debris on lower shore. 
RECORD. Komimbo Bay - i. 

NOTES. The specimen is complete with 19 setigers plus one or two posterior 
achaetous segments. A cephalic plaque is present, the rim of which has six well 
defined lobes along the dorsal margin. There are a few prostomial pigment spots. 
Setigers i to 3 each have a single large aciculum. A caudal funnel is present and 
this carries a median ventral cirrus that is more than 1-5 times the length of the other 
cirri. Questionably identified as Praxillella kollikeri (Mclntosh, 1885, as Pr axilla 
kollikeri). 



Praxillella sp. 

HABITAT. Silty fine coral sand at LWM. 
RECORD. Matiu Is. - i. 

NOTES. The complete specimen has 19 setigers plus two posterior achaetous 
segments. A cephalic plaque is developed with nuchal organs extending over nine- 
tenths of its length. The cephalic rim has posterior clefts only. Pigment spots on 
the prostomium are profuse. There are three or four uncini on setigers i and 2 and 
four on setiger 3. A caudal funnel is not present. The anal plug is just protruding 
and is surrounded by cirri of equal length. The specimen resembles P. affinis (Sars) 
but is probably a new species. 



Euclymeninae sp. A 

HABITAT. Muddy sand at 9 m depth. 
RECORD. ML 56-1. 

NOTES. This specimen is an anterior fragment of nine setigers. The cephalic 
plaque has nuchal organs extending across three-quarters of its length. Prostomial 
pigment spots are absent. The first setiger is not clearly separated from the 
prostomium. One pair of acicula is present on setiger i, two pairs on setiger 2 and 
one or two pairs on setiger 3, with uncini on the following segments. 



Euclymeninae sp. B 

HABITAT. Between boulders in muddy silt at MTL. 
RECORD. Tetel Is. - i. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 189 

NOTES. This incomplete specimen is composed of the prostomium and eight 
setigers. The nuchal organs extend for nine-tenths of the length of the cephalic 
plaque and there are very shallow clefts around the medial and posterior margins of 
the cephalic rim. Pigment spots are present ventrally. Setiger i has one pair of 
acicula and setigers 2 and 3 have two pairs, the subsequent setigers having uncini. 



Euclymeninae sp. C 

HABITAT. Silty sand at 24 m depth. 
RECORD. ML 230-1. 

NOTES. The specimen is fragmentary, consisting of 15 anterior setigers. The 
cephalic plaque is unusual in having pigment spots within the plaque as well as on the 
ventral surface. The cephalic rim is very high and has posterior clefts only. 



Subfamily NICOMACHINAE 
? Nicomache sp. 

HABITAT. Coarse coral sand below LWM. 
RECORDS. Tetel Is. - 2. 

NOTES. The material consists of two large anterior fragments with seven and 
eight setigers. They appear to be identical except that one specimen has only large 
setae on setigers I to 3 but the other specimen has four small setae on setiger i, a 
small (left) and a large (right) seta on setiger 2 and one large seta on either side of 
setiger 3. The latter specimen may be abnormal due to aberrant growth or re- 
generation. In both specimens the first three setigers are light, the rest being grey- 
brown to dark in colour. 



Family OWENIIDAE 

Myriochele eurystoma Caullery, 1944 

(Fig. 15. A) 

Myriochele eurystoma Caullery, 1944 : 52, fig. 42. a-d, f. 

HABITAT. Silty sand at 35 m depth. 
RECORD. ML 72-1. 

NOTES. The specimen is an anterior fragment about 15 mm long and has a 
diameter of 0-5 mm. It has a large mouth with a prominent buccal organ on its 



i go 



P. E. GIBBS 



ventral margin (fig. 15. A). The prostomium and anterior segments are pigmented 
reddish-brown, with two eye-spots situated ventro-laterally. Between the latter, 
running over the dorsal surface, there is a thin unpigmented band. The tube is 
composed chiefly of sponge spicules cemented with their long axes around the 
circumference of the tube, the latter having a diameter of about 1-2 mm. 

The specimen is referred to M. eurystoma on account of its large mouth and its 
pigmentation. Caullery notes that a specimen from 32 m depth was pigmented but 
specimens from deeper waters (131-1570 m) lacked colour. The details of the 
posterior region are unknown. 

DISTRIBUTION. East Indies. 



Myriochele heruensis n. sp. 

(Fig. 15. B-G) 

DESCRIPTION. Of the fourteen specimens taken at ML 134, only three could 
be removed intact from their tubes. The holotype is 7-6 mm long for 18 segments 
and the two other complete paratypes measure 6-6 and 7-5 mm for 16 and 17 seg- 
ments respectively. The largest specimen probably measured between 10 and 12 
mm long when intact. The diameter of the worm is between 0-28 and 0-40 mm. 






H G F E 

FIG. 15. Myriochele eurystoma. (A) Anterior region. Myriochele heruensis n.sp. (B-D) 
Ventral and lateral views of anterior region. (E) Dorsal view of posterior region. (F-G) 
Profile and face views of uncinus. Myriochele sp. (H) Ventral view of anterior region. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 191 

The prostomium is short with a rounded anterior margin (fig. 15. B). In those 
specimens preserved with the prostomium protracted, a shallow groove can be 
discerned extending across the dorsal surface of the prostomium, terminating on 
either side at the level of ventro-lateral eye-spots (fig. 15. c). In other specimens 
the prostomium is contracted, and in these the dorsal groove appears as a distinct 
fold (fig. 15. D). This groove or fold is quite marked in all specimens and serves as 
the characteristic feature of the species. In addition to the two red eye-spots, two 
small patches of red pigment are present on the antero-dorsal surface of the prosto- 
mium and there is also a thin red line along the posterior margin of the dorsal groove. 

The first three setigers are short. From setiger 4 there is a progressive increase 
in the length of the segments, reaching a maximum in the middle body region, and 
then successive segments become shorter. The pygidium has two short, stout anal 
cirri (fig. 15. E). 

The setae are of the usual Myriochele types. Notosetae are slender capillaries 
with short lateral processes along either side of the blade. Neurosetae, present from 
setiger 4, are uncini, each with two subequal teeth at the apex (fig. 15. F-G). In 
the middle body segments, the tori are composed of 7 or 8 vertical rows of uncini. 

The tube is composed of shell fragments which are cemented with their flat 
surfaces at right angles to the long axis of the tube. The latter has a diameter of 
0-8 to i-o mm and a maximum length of about 25 mm. 

As noted above the distinct groove across the dorsal surface of the prostomium 
is sufficient to distinguish M. heruensis from all other species. This feature may be 
interpreted as a feeding adaptation in that it provides a mechanism whereby the 
position of the mouth can be changed. 

HABITAT. Sand at 16 m depth. 

RECORDS. ML 134 - 14. 

British Museum (Natural History) Registration No. Holotype 1970-77 

Paratypes 1970-78 



Myriochele sp. 

(Fig. 15. H) 

HABITAT. Sand at 5 m depth. 
RECORD. ML 229 - i. 

NOTES. The specimen lacks posterior segments but probably measured about 
12 mm long when intact. It has a diameter of 0-24 mm. The anterior margin of the 
prostomium is bluntly rounded and, ventrally, the mouth extends about half the 
prostomial length, ending at the level of two red eye-spots which are ventro-lateral 
in position (fig. 15. H). 



192 P. E. GIBBS 

The tube has a diameter of 0-4 mm and consists of sand and shell fragments 
cemented with their long axes along the length of the tube. 

The specimen shows a general similarity to M. minor Caullery but differs in having 
eye-spots. 



Owenia fusiformis Delle Chiaje, 1844 

Owenia fusiformis : Fauvel, 1927 : 203, fig. 71. a-f; 1953 : 39 1 ' fig- 2O 3- a ~f; Day, 1967 : 649, fig. 
31.1. e-j. 

HABITAT. Medium and coarse sand, 2-18 m depth. 

RECORDS. ML 96-3; ML 118 - 3; ML 134- 5; ML 190 - 2; ML 192 - 13; 
ML 203 - i ; ML 204 - 60 ; ML 229 - 2 ; ML 283 - i. 

DISTRIBUTION. Cosmopolitan. 



Family SABELLARIIDAE 
Lygdamis ehlersi (Caullery, 1913) 

Tetreres ehlersi Caullery, 1913 : 201, figs a-d. 
Lygdamis ehlersi: Caullery, 1944 : 62, figs 49-50. 

HABITAT. Silty sand at LWM. 
RECORDS. Fintry Pt. - 5. 

NOTES. The largest specimen is 50 mm long for 50 segments. The tube is 
constructed of gravel, shell and calcareous alga fragments. The chief characters 
include the following: on the operculum, there are 19 or 20 external and 6 or 7 
internal paleae on each side and 10 groups of buccal cirri. The median cirrus (lobe 
Preoral) is stout, longer than the palps, and has a truncated tip which is darkly 
pigmented a bluish-black colour. There are 13 to 15 pairs of branchiae. 

Comparing these specimens with the type description they have fewer external 
(20 compared to 25) and internal (6 or 7 instead of 12) paleae but correspond in all 
other details. 

Although Hartman (1959) indicates that L. ehlersi may be a synonym of L. indicus 
the two species appear to be distinct and separable on the form of the median cirrus. 
In L. ehlersi the median cirrus is stout, longer than the palps and has a truncated 
tip while in L. indicus this organ is shorter than the palps and tapers to a point. 

DISTRIBUTION. East Indies. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 193 

Lygdamis indicus Kinberg, 1867 

Lygdamis indicus: Johansson, 1925 : 8, fig. 2. 2-7; Fauvel, 1953 : 399, fig. 209. a-k; Day, 1967 : 
677. fig- 33-3- c-h. 

HABITAT. Undersurface of coral boulder at LWM. 
RECORD. Pirikale Is. - i. 

NOTES. The specimen is 30 mm long (diameter = 4 mm) with 48 setigers. It 
has 37 external and 17 internal paleae on each side. The tube is a fragile structure 
constructed of shell fragments. 

DISTRIBUTION. Tropical Indo-west-Pacific. 



Family PECTINARIIDAE 
Pectinaria (Pectinaria) antipoda Schmarda, 1861 

Pectinaria (Pectinaria) antipoda Schmarda, 1861 : 46, pi. 24, fig. 199, with text-figs; Pruvot, 
1930 : 78, pi. 3, figs 93-95; Fauvel, 1953 : 403, fig. 211. e-g. 

HABITAT. Mud and sand, 4-22 m depth. 
RECORDS. ML 37 - i ; ML 203 - i ; ML 229 - i. 

NOTES. All three specimens are too small (less than 10 mm) to be certain of their 
identity but they appear to belong to this Indo-west-Pacific species. 

DISTRIBUTION. Persian Gulf; West Pacific. 



Family AMPHARETIDAE 

Subfamily MELINNINAE 
Isolda pulchella Miiller, 1858 

Isolda sibogae Caullery, 1944 : 102, fig. 83. a-h. 

Isolda pulchella: Day, 1963 : 434 (synonymy); 1967 : 691, fig. 35.1. k-n. 

HABITAT. Mud and silty sand, 2-18 m depth. 
RECORDS. ML 56- i; ML 196- i; ML 218- i. 
DISTRIBUTION. Atlantic and Indian Oceans; East Indies. 



194 



P. E. GIBBS 



Subfamily AMPHARETINAE 
? Sosane wireni Caullery, 1944 

(Fig. 16. A-B) 

Sosane wireni Caullery, 1944 : 87, fig. 69. a-e. 
HABITAT. Silty sand at 24 m depth. 
RECORD. ML 230 - i. 

NOTES. The specimen is 7 mm long for 26 setigers of which 15 are thoracic and 
ii abdominal. It is referred to S. wireni on account of the branchiae being arranged 
with three pairs in a transverse line across setiger i and a smaller fourth pair on 
setiger 2. However there is some doubt as to the identity since Caullery omits any 
reference to the diagnostic features of the genus, namely, the presence of (i) a 
nephridial papilla on the branchial ridge and (ii) specialised notosetae on elevated 
posterior notopodia (Day, 1964). There does not appear to be a nephridial papilla 
present on the Solomon Islands specimen but the notopodia of setiger 13 are elevated 
(fig. 16. A) and each carries about 20 modified notosetae which have a 'penicillate' 
appearance (fig. 16. B). 

DISTRIBUTION. East Indies. 





B 



FIG. 16. ? Sosane wireni. (A) Elevated notopodium of setiger 13. 
(B) Modified notosetae from same setiger. 



Family TEREBELLIDAE 

Subfamily TRICHOBRANCHINAE 

Terebellides stroemi Sars, 1835 

Terebellides stroemi: Fauvel, 1927 : 291, fig. 100. i-q; 1953 : 436, fig. 231. i-q; Day, 1967 : 713, 
fig. 36.1. f-j. 

HABITAT. Mud and silty sand, 2-24 m depth. 

RECORDS. ML 155 - 31 ; ML 157 - 3 ; ML 196 - i ; ML 218 - i ; ML 230 - 3. 

DISTRIBUTION. Cosmopolitan. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 195 

Subfamily POLYGIRRINAE 
Amaeana trilobata (Sars, 1863) 

Amaea trilobata: Fauvel, 1927 : 285, fig. 99. a-e. 
Amaeana trilobata: Day 1967 : 718, fig. 36.3. e-h. 

HABITAT. Mud, 11-22 m depth. 

RECORDS. ML 29 - i ; ML 194 - i. 

DISTRIBUTION. North Atlantic; Mediterranean; southern Africa; Japan. 

Lysilla ubianensis Caullery, 1944 

Lysilla ubianensis Caullery, 1944 : 197, fig. 156. a-e; Day, 1967 : 721, fig. 36.3. i-j. 

HABITAT. Silty mud, MTL to LWM, and at 13 m depth 
RECORDS. Tetel Is. - 3 ; Komimbo Bay - i ; ML 69-1. 
DISTRIBUTION. East Africa; East Indies. 

Subfamily THELEPINAE 
Euthelepus kinsemboensis Augener, 1918 

Euthelepus kinsemboensis Augener, 1918 : 548, pi. 6, fig. 161, pi. 7, fig. 250, text-fig. 95; Fauvel, 
I 93 0: 553. fig- 9- a-f; 194? : 7 8 - fi g- 75- a'-d'; Day, 1967 : 726, fig. 36.5. e-i. 

HABITAT. Under coral boulder on reef platform. 
RECORD. Graham Pt. - i. 

NOTES. The specimen is an anterior fragment of 34 setigers which corresponds 
in detail to the specimen from New Caledonia described by Fauvel except that the 
branchial filaments, situated on segments 2, 3 and 4, number 10, 8 and 6 respectively, 
not 12, 8 and 4. The type specimen from Angola has only 12 branchial filaments 
instead of 24, but according to Fauvel (1930) this variation is attributable to the 
difference in the size of the specimens. 

DISTRIBUTION. Angola; New Caledonia. 

Subfamily TEREBELLINAE 
Eupolymnia nebulosa (Montagu, 1818) 

Polymnia nebulosa: Fauvel, 1927 : 257, fig. 89. a-g; 1953 : 419, fig. 219. a-g. 
Eupolymnia nebulosa: Imajima & Hartman, 1964 : 337; Day, 1967 : 744, fig. 36.9. f-h. 

HABITAT. Under coral boulder on silty sand at LWM. 

RECORD. Graham Pt. - i. 

DISTRIBUTION. Atlantic; South Africa; Mediterranean; Indo-west Pacific. 



ig6 P. E. GIBBS 

Loimia medusa (Savigny, 1820) 
Loimia medusa: Fauvel, 1953 : 4 J 6, fig- 218. a-f; Day, 1967 : 743, fig. 36.9. a-e, 

HABITAT. A wide variety of deposits, silty mud to Acropora rubble, from above 
MTL to LWM on reef platforms; in crevices within reef platforms and amongst 
encrustations on wharf piles ; silty sand, 9-24 m depth. 

RECORDS. Tetel Is. - 3 ; Komimbo Bay - 9 ; Mamara Pt. - i ; Matiu Is. - i ; 
Lauvie Is. - 6 ; Fintry Pt. - 9 ; Yandina - 2 ; Maramasike Pg. - i ; ML 69 - i ; 
ML 96 - i ; ML 230 - i. 

NOTES. The largest specimens, up to 40 cm in length with over 200 segments, 
were found in coral debris deposits (chiefly Acropora fragments) on reef platforms. 
Their tubes were loosely constructed of Halimeda, coral and shell fragments and 
extended to a depth of over 30 cm. Although conspicuous on account of their 
white feeding tentacles spread over the surface, specimens are difficult to capture 
in this habitat because of their ability to rapidly withdraw to the bottom of their 
deep tubes when disturbed. However smaller specimens which are commonly 
found under coral boulders lying in silty mud are easily taken. 

DISTRIBUTION. Cosmopolitan in temperate and tropical waters. 

Pista dibranchis n. sp. 

(Fig. 17. A-E) 

DESCRIPTION. A specimen from Tetel Island has been selected for the holotype. 
It is incomplete and measures 12 mm for 32 segments. The largest of the paratypes 
is 40 mm long for 130 segments. The body diameter is about i-o mm. 

The tentacular lobe bears numerous tentacles and is without eye-spots. Semi- 
circular lateral lobes are present on segments 2 and 3. A pair of stalked branchiae 
arises from the anterior margin of the second segment and each has numerous 
filaments arranged spirally, with a maximum of 6 or 7 whorls, around the central 
stalk (fig. 17. A) ; often one of the pair is much larger than the other. In larger 
specimens the nephridial papillae on segments 6 and 7 (setigers 3 and 4) are cons- 
picuous. 

Notosetae commence on segment 4 and extend to segment 20 (setiger 17). Each 
bundle consists of broad-winged, smooth-tipped capillaries of two types, namely, a 
longer stout-shafted form (fig. 17. B) and a shorter, slender form with a recurved tip 
(fig. 17. c). There are usually about n of each type in a bundle. Uncini start on 
segment 5 (setiger 2) and are arranged in single rows on the anterior segments, 
becoming alternate in the posterior thorax. Uncini are of similar form throughout 
the thorax and each has 5 to 7 teeth surmounted by 15 to 20 smaller teeth above the 
main fang (fig. 17. D, E). The dental formula is thus: MF : 5-7 : 15-20. The uncini 
lack basal prolongations or shafts, even in the first row. Abdominal uncini are 
borne on short, square pinnules. 

The tube is composed of sand grains cemented together with mud particles. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 



197 



At first it was assumed that these specimens possessing only one pair of branchiae 
were P. typha juveniles in which the second pair of branchiae was either undeveloped 
or had been lost. However subsequent examination revealed that the uncini of the 
anterior segments lacked the basal shafts found in P. typha, and also some of the 
specimens contain coelomic oocytes, apparently approaching a mature condition. 

In having only one pair of branchiae, P. dibranchis resembles P. vinogradovi 
Uschakov but lacks the transverse dorsal membrane between the branchial stalks 
that is present in the latter species. According to Uschakov (1955), P. cristata 
(Miiller) may also have one pair of branchiae but this species differs in possessing 
shafted uncini. In terms of the structure of the branchial apparatus, P. dibranchis 
is intermediate between P. unibranchia Day, which has a single branchia, and P. 
typha Grube, which has two pairs, all three species having branchiae with spirally 
arranged, or whorled, filaments. 

HABITAT. A wide variety of deposits, silty mud to coarse sand and shell gravel, 
from MTL to 24 m depth. 

RECORDS. Haroro - i ; Tetel Is. - 4 (including holotype) ; Graham Pt. - i ; 
Fintry Pt. - 3 ; ML 38 - i ; ML 204 - i ; ML 230 - i ; ML 283 - 6; ML 296 - i. 

British Museum (Natural History) Registration No. Holotype 197079 

Paratypes 1970-80-89 

NOTE. The habitat of P. dibranchis overlaps that of P. typha ; on Tetel Island the 
two species were taken from the same silty mud deposit on the reef platform at about 
MTL. 






FIG. 17. Pista dibranchis n.sp. (A) Dorsal view of anterior region. 
(B-C) Thoracic notosetae. (D-E) Profile and face views of uncinus. 



ig8 P. E. GIBBS 

Pista typha (Grube, 1878) 

Terebella (Pista) typha Grube, 1878 : 232, pi. 12, fig. 4. 
Pista typha: Fauvel, 1953 : 424, fig. 222. a-c. 

HABITAT. Silty sand and mud, MTL to LWM. 
RECORDS. Tetel Is. - 23; Fintry Pt. - i. 
DISTRIBUTION. Indo-west-Pacific. 



Reteterebella queenslandia Hartman, 1963 

Reteterebella queenslandia Hartman, 1963 : 355, figs 1-3. 

HABITAT. Acropora rubble at LWM. 
RECORDS. Tetel Is. - i ; Maraunibina Is. - i. 

NOTES. The genus Reteterebella Hartman differs from the closely allied genus 
Eupolymnia Verrill in lacking notosetae on segment 4, thus having 16 instead of 17 
thoracic setigers, and also in the first appearance of the double rows of uncini on the 
sixth (segment 10 or setiger 6), not the seventh uncinigerous segment (= setiger 8 in 
Eupolymnia). 

DISTRIBUTION. North-east Australia (Queensland). 



Terebella ehrenbergi Grube, 1870 

Terebella ehrenbergi: Fauvel, 1953 : 421, fig. 220. a-c; Day, 1967 : 748, fig. 36.10. g-i. 

HABITAT. Crevices in reef platform. 

RECORDS. Matiu Is. - 2. 

DISTRIBUTION. Tropical Indo-west-Pacific. . 



Family SABELLIDAE 
Branchiomma cingulata (Grube, 1870) 

Dasychone cingulata: Willey, 1905 : 308, pi. 7, figs 170-173; Fauvel, 1953 : 442, fig. 234. f-h. 
Branchiomma cingulata: Johansson, 1927 : 161, text-fig. 14.1-2; Imajima & Hartman, 1964 : 355. 

HABITAT. Embedded in Acropora at LWM. 
RECORD. Tetel Is. - i. 

NOTES. The specimen is small, about 7 mm long. It has six radioles on each 
side of the branchial crown and the thoracic region consists of five segments. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 199 

Specimens from Ceylon and Japan have a greater number of radioles and eight 
thoracic segments but are larger-sized, which may account for these differences. 
DISTRIBUTION. Indo-Pacific. 



Hypsicomus phaeotaenia (Schmarda, 1861) 

Hypsicomus phaeotaenia : Fauvel, 1953 : 447, fig. 236. a-1; Day, 1967 : 761, fig. 37.2. i-n. 
HABITAT. Embedded (boring?) in coral, especially Forties, and in beachrock. 

RECORDS. Kokomtambu Is. - 1 ; Tetel Is. - 4 ; Komimbo Bay - 3 ; Mamara Pt. - 1 ; 
Matiu Is. - 5 ; Paleki Is. - i ; Lauvie Is. - 7; Graham Pt. - 3. 

DISTRIBUTION. Western Africa; Mediterranean; Indo-west-Pacific. 



Megalomma intermedium (Beddard, i 

Branchiomma intermedium Beddard, 1888 : 261, pi. 21, figs 4-7; Fauvel, 1953 : 444, fig. 234. e. 
Megalomma intermedium: Pillai, 1965 : 164. 

HABITAT. Embedded in Porites boulder towards LWM. 

RECORD. Tetel Is. - i. 

DISTRIBUTION. Mergui Archipelago; Philippine Islands. 



Megalomma linaresi (Rioja, 1918) 

Branchiomma linaresi: Fauvel, 1927 : 317, fig. no. a-1. 

HABITAT. Embedded in Acropora at LWM and at 5 m depth. 

RECORDS. Tetel Is. - i ; Yandina - i. 

NOTES. The larger specimen is 45 mm long, including the branchial crown which 
has a length of 10 mm. There are 16 to 19 radioles on each side. The dorsal radioles 
carry the largest subterminal eyes, which gradually diminish in size on the lateral 
radioles and are absent from the ventral ones. The collar gapes widely on the 
dorsal side: it has lateral notches and large ventral lobes. There are 8 thoracic 
segments, the notosetae of which consist of narrow-winged capillaries and sub- 
spatulate setae. 

M. linaresi is known from northern Spain and the Mediterranean. Despite the 
absence of records from the Indian Ocean, the specimens from the Solomon Islands 
appear to be identical. 

DISTRIBUTION. North Atlantic (Spain) ; Mediterranean. 



200 P. E. GIBBS 

Megalomma quadrioculatum (Willey, 1905) 

Branchiomma quadrioculatum Willey, 1905 : 307, pi. 7, figs 168-169. 
Megalomma quadrioculatum: Day, 1967 : 758, fig. 37.1. h-o. 

HABITAT. Amongst Phyllochaetopterus socialis tubes on reef platform towards 
LWM ; amongst serpulid tubes on undersurface of coral boulder. 

RECORDS. Cape Esperance - 36; Komimbo Bay - i. 

NOTES. The specimens were removed from sandy tubes constructed amongst 
the sponges and other encrusting organisms around the bases of the chaetopterid 
tubes. The larger specimens measure 13 to 14 mm long. The thorax consists of 
6 or 7 segments with notosetae of two types, namely narrow-winged capillaries and 
paleae. There are 6 to 9 radioles on each side of the branchial crown and, in most 
specimens, only the two dorsal radioles carry subterminal eyes. However, a few 
specimens have smaller eyes on one or two radioles besides the dorsal radioles. 

The type specimen from Ceylon (23-5 mm in length) is almost twice the size of the 
Solomon Islands specimens and has 14 radioles per branchial lobe with subterminal 
eyes on the four dorsal radioles. It would seem therefore that the second (and 
occasionally a third) pair of eyes develops at a later stage and, until acquired, M. 
quadrioculatum bears a superficial resemblance to M . bioculatum (Ehlers) from which 
it can be distinguished by the presence of paleae in the notopodia. 

DISTRIBUTION. Indian Ocean. 

Megalomma trioculatum Reish, 1968 

Megalomma trioculatum Reish, 1968 : 226, fig. 5. i-io. 

HABITAT. Sand, 2-9 m depth. 
RECORDS. ML 96 - i; ML 296- i. 

NOTES. The two specimens measure 15 to 16 mm long, the branchial crown 
being 2-0 to 2-5 mm in length. Both have 6 thoracic setigers (instead of 8 or 9) and 
the collar gapes widely on the dorsal side of the first setiger instead of extending to 
the mid-segmental line. Otherwise they agree closely with type description. 

At the bases of the two dorsal radioles, there are longitudinal streaks of an orange- 
brown pigment and the radioles are banded a similar colour. The subterminal eyes 
are brown or mauve. The tube is a fragile structure coated with sand grains. 

DISTRIBUTION. Marshall Islands. 

Megalomma vesiculosum (Montagu, 1815) 

Branchiomma vesiculosum: Fauvel, 1927 : 315, fig. 109. a-q. 
Megalomma vesiculosum: Day, 1967 : 758, fig. 37.1. p-u. 

HABITAT. Muddy sand between MTL and LWM. 
RECORDS. Tetel Is. - 20. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 201 

NOTES. The subterminal eyes are entire, not divided as in M. pacificum Johansson 
although the latter may be a synonym of M. vesiculosum, according to Mesnil & 
Fauvel (1939). 

DISTRIBUTION. Atlantic Ocean; Mediterranean; Indian Ocean. 



Potamilla ehlersi Gravier, 1906 

Potamilla ehlersi Gravier, igo6a : 37; 1908 : 87, pi. 6, fig. 260-264; Fauvel, 1953 : 449, 
fig. 238. g-i. 

HABITAT. Amongst Phyllochaetopterus socialis tubes on reef platform towards 
LWM. 

RECORD. Cape Esperance- i. 

NOTES. The specimen is 13 mm long (branchial crown = 3 mm). There are 
nine radioles on each branchial lobe ; the dorsal ones are without eyes but these are 
present on the next 4 or 5 radioles. There are n segments in the thorax, the 
anterior half of which is coloured chocolate-brown on the dorsal side. The tube is 
membraneous, covered with sand grains, and, before dissection, its aperture was 
rolled, as found in P. reniformis (Muller) (cf. Fauvel, 1927, fig. 107 1). 

DISTRIBUTION. Tropical Indo-west-Pacific. 



Sabellafusca Grube, 1870 

Sabella fusca: Fauvel, 1927 : 302, fig. 104. a-f; Day, 1967 : 764, fig. 37.2. t-v. 
HABITAT. Beachrock. 
RECORDS. Lauvie Is. - 3. 
DISTRIBUTION. Mediterranean; tropical Indo-west-Pacific. 



Sabella melanostigma Schmarda, 1861 

Sabella melanostigma Schmarda, 1861 : 36, pi. 22, fig. 190, with text-figs; Fauvel, 1953 : 439, 
fig. 232. h-n. 

HABITAT. Mud and silty sand, 24-33 m depth. 
RECORDS. ML 100 - i ; ML 230 - 5. 

NOTES. The specimens are only about 12 mm long and were extracted from 
narrow tubes (0-65 mm diameter) composed of mud particles. They are referred to 



P. E. GIBBS 



this species because they possess the peculiar 'shovel-like' pick-axe setae. However 
pigment spots above the parapodia are lacking. There are 7 radioles, each with 2 
or 3 pairs of eye-spots, on either branchial lobe. The ventral lobes of the collar are 
large and there is a wide gap between the dorsal lobes. 

DISTRIBUTION. Circumtropical. 



Sabellastarte sanctijosephi (Gravier, 1906) 

Eurato sancti-josephi Gravier, igo6a : 42; 1908 : 105, pi. 7, figs 281-283, pi. 8, figs 284-285. 
Sabellastarte indica: Fauvel, 1953 : 445, fig. 235. a-h. 
Sabellastarte sanctijosephi: Day, 1967 : 771, fig. 37.5. f-i. 

HABITAT. In crevices in Forties boulders at LWM (Gibbs, 1969, fig. 137). 

RECORDS. Tetel Is. - i ; Paleki Is. - 5. 

DISTRIBUTION. Western Africa; tropical Indo-west-Pacific. 



Family SERPULIDAE 

Subfamily SERPULINAE 

Hydroides minax (Grube, 1878) 

Serpula minax Grube, 1878 : 269, pi. 15, fig. 5. 

Hydroides minax: Fauvel, 1953 : 4^o, fig. 241. f; Pillai, 1960 : 8, text-fig. 3. a-e. 

HABITAT. Amongst Phyllbchaetopterus socialis tubes at LWM. 
RECORDS. Matiu Is. - 2. 

NOTES. These specimens were referred to H. monoceros Gravier, a closely allied 
species, in an earlier paper (Gibbs, 1969). 
DISTRIBUTION. Ceylon; Philippine Islands. 



Hydroides uncinata (Philippi, 1844) 

Hydroides uncinata: Fauvel, 1927 : 357, fig. 122. a-h; Day, 1967 : 805, fig. 38.4. h-i. 
HABITAT. On Acropora at 5 m depth. 
RECORD. Yandina-i. 
DISTRIBUTION. Atlantic Ocean; Mediterranean; East Africa; Japan. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 203 

Mercierella enigmatica Fauvel, 1923 

Mercierella enigmatica: Fauvel, 1927 : 360, fig. 123. a-o; Day, 1967 : 812, fig. 38.5. o-s. 
Neopomatus uschakovi Pillai, 1960 : 28, pi. i, figs 1-2; text-figs 10. h, n. a-h, 12. a-h; Straughan, 
1966 : 139 (synonymy). 

HABITAT. Brackish water - on stones and in sponge. 

RECORDS. Komimbo Bay (freshwater creek) - num. ; mouth of Lunga River at 
Lunga Pt. - 28. 

NOTES. In a study of the brackish water serpulids along the east coast of 
Australia, Straughan (1966) discovered that there is a continuous cline in the characters 
used to separate the two genera Mercierella Fauvel and Neopomatus Pillai. The 
Mercierella form is characteristic of the southern populations around Sydney and 
the warm water Neopomatus form is typical for the Queensland populations. 

The material from Guadalcanal is typical Neopomatus in that there are one or 
two rows of outwardly directed spines on the operculum, the collar and thoracic 
membranes are fused dorsally for up to three-quarters of the length of the thorax, 
and the collar setae have modified proximal teeth. 

DISTRIBUTION. Cosmopolitan in estuarine water. 



Serpula hartmanae Reish, 1968 

Serpula hartmanae Reish, 1968 : 228, fig. 5. 11-16. 

HABITAT. Attached to coral on reef platform. 
RECORDS. Mamara Pt. - i ; Komimbo Bay - 3. 
DISTRIBUTION. Marshall Islands. 



Serpula vermicularis Linnaeus, 1767 

Serpula vermicularis: Fauvel, 1927 : 351, fig. 120. a-q; 1953 : 454, fig. 239. a-q; Dew, 1959 : 22, 
fig. 3. a-h; Day, 1967 : 809, fig. 38.5. a-h. 

HABITAT. Attached to coral boulders on reef platform. 
RECORDS. Tetel Is. - 2 ; Komimbo Bay - 5. 
DISTRIBUTION. Cosmopolitan. 



Spirobranchus giganteus (Pallas, 1766) 

Spirobranchus giganteus : Fauvel, 1953 : 462, fig. 242. a-g; Day, 1967 : 803, fig. 38.3. h-k. 

HABITAT. Embedded in living Porites boulders (Gibbs, 1969, fig. 138) and 
encrusting on reef platform. 



204 P. E. GIBBS 

RECORDS. Kokomtambu Is. -2; Tetel Is. - v. num.; Komimbo Bay -num.; 
Mamara Pt. - i ; Matiu Is. - 9; Paleki Is. - v. num. ; Sifola - i ; Graham Pt. - num. 

NOTES. This species is common in moderately sheltered conditions and the 
largest specimens, over no mm in length and 10 mm in diameter, were extracted 
from living Porites at Paleki Island. In comparison, encrusting forms on dead 
coral are much smaller in size. 

DISTRIBUTION. West Indies; Indo-west-Pacific. 



Spirobranchus coutierei (Gravier, 1908) 

Pomatoceropsis coutierei Gravier, 1908 : 125, pi. 8, figs 294-299, text-figs 482-487. 

HABITAT. Embedded in Porites towards LWM. 
RECORDS. Tetel Is. - 3. 

NOTES. Following Straughan's (1967) key to the Indo-west-Pacific species, this 
identification is based on the following two characters - (i) the operculum has three 
processes which are deeply divided so as to give six distinct horns, and (ii) the 
interbranchial membrane is fimbriated. 

DISTRIBUTION. East Africa; northern Australia (Straughan, 1967^). 
Vermiliopsis glandigerus Gravier, 1908 

Vermiliopsis glandigerus Gravier, 1908 : 121, pi. 8, figs 290-291, text-figs 476-481; Fauvel, 
1953 : 467, fig. 242. k; Day, 1967 : 813, fig. 38.6. g-i. 

HABITAT. Encrusting on reef platform. 

RECORD. Matiu Is. - i. 

DISTRIBUTION. Western and southern Africa; Indo-west-Pacific. 

Subfamily FILOGRANINAE 
Filograna implexa Berkeley, 1835 

Salmacina dysteri: Fauvel, 1927 : 377, fig. 129. c-k; 1953 : 476, fig. 250. c-k; Dew, 1959 : 50, 

fig. 19. a-g. 
Filograna implexa: Fauvel, 1927 : 376, fig. 129. a-b; Day, 1955 : 450 (synonymy); 1967 : 817, 

fig. 38.7. a-h. 

HABITAT. Encrusting on an oyster shell attached to wharf pile. 
RECORDS. Yandina - num. 

NOTE. No individual with an operculum developed at the end of a branchial 
filament was noted amongst the colony; the specimens thus correspond to the 
Salmacina form. 

DISTRIBUTION. Cosmopolitan. 



THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 205 

ADDENDUM 

Recently Professor J. E. Morton forwarded a small collection of polychaetes which 
he collected in the Solomon Islands during the 1965 Expedition. This material 
includes specimens of a further two species, bringing the total number of recorded 
species to 222. 

The species are the following : 



Family NEREID AE 
Perinereis neocaledonica Pruvot, 1930 

Perinereis neocaledonica Pruvot, 1930 : 50, pi. 3, figs 77-79, text-fig. 4. a-c; Fauvel, 1932 : 107; 
1953 : 2ii, fig. 108. c-g; Tampi & Rangarajan, 1964 : 106. 

RECORDS. Batuona Is - 10. 
DISTRIBUTION. Tropical Indo- west-Pacific. 



Family GLYCERIDAE 
Glycera subaenea Grube, 1878 

Glycera subaenea Grube, 1878 : 184, pi. 8, fig. 8; Fauvel, igiga : 425, pi. 16, figs 48-51; Hartman 
& Imajima, 1964 : 164; Day, 1967 : 363, fig. 16.3. k-n. 

RECORDS. Komimbo Bay - 3. 
DISTRIBUTION. Indo-west-Pacific. 



REFERENCES 

ACHARI, G. P. K. 1968. Studies on new or little known polychaetes from Indian Seas. 

i. Trochochaeta watsoni (Fauvel) and Poecilochaetus serpens Allen. /. mar. biol. Ass. 

India 10 : 99-106. 

ALLEN, E. J. 1904. The anatomy of Poecilochaetus. Q. Jl microsc. Sci., n.s. 48 : 79-151. 
AUGENER, H. 1913. Polychaeta Errantia. Fauna Sudwest-Aust. 4 : 65-304. 

1914- Polychaeta Sedentaria. Fauna Sudwest-Aust. 5 : 1-170. 

1918. Polychaeta. Beitr. Kennt. Meeresfauna Westafr. 2 : 67-625. 

1922. Results of Dr. E. Mjosberg's Swedish Scientific Expeditions to Australia 1910- 

1913. 32. Polychaeten. K. svenska Vetensk. Akad. Handl. 63 (6) : 1-49. 

- 1924. Papers from Dr. Th. Mortensen's Pacific Expedition 1914-1916. 1 8. Polychaeta 2. 
Polychaeten von Neuseeland. i. Errantia. Vidensk. Meddr dansk naturh. Foren. 75 : 
241-441. 

1926. Papers from Dr. Th. Mortensen's Pacific Expedition 1914-1916. 34. Polychaeta 3. 
Polychaeten von Neuseeland. 2. Sedentaria. Vidensk. Meddr dansk naturh. Foren. 

81 : 157-294- 

1927. Papers from Dr. Th. Mortensen's Pacific Expedition 1914-1916. 38. Polychaeten 
von Sudost-und Siid-Australien. Vidensk. Meddr dansk naturh. Foren. 83 : 71-275. 



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AUGENER, H. 19273. Polychaeten von Neu-Pommern. Sber. Ges. naturf. Freunde, Berl. vol 

for 1926 : 119-152. 
BANSE, K. 1959. On marine Polychaeta from Mandapam (South India). J. mar. biol. Ass. 

India 1 : 165-177. 
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trustees of the Indian Museum, Calcutta, by Dr. John Anderson, F.R.S. /. Linn. Soc. 

(Zool.) 21 : 256-266. 
BHAUD, M. 1969. Remarques syst6matiques et biogeographiques sur le genre Mesochaetopterus 

Potts, 1914. Vie Milieu B 20 : 325-332. 
BURROWS, W. 1945. Periodic spawning of 'palolo' worms in Pacific waters. Nature, Lond. 

155 : 47-48. 

1955- 'Palolo', notes on the periodic appearance of the annelid worm Eunice viridis (Gray) 

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THE POLYCHAETE FAUNA OF THE SOLOMON ISLANDS 211 

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Dr. P. E. GIBBS 

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(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 21, No. 6 of the Zoological series. 
The abbreviated titles of periodicals cited follow those 
of the World List of Scientific Periodicals. 



World List abbreviation 
Bull. Br. Mus. nat. Hist. (Zool.) 



Trustees of the British Museum (Natural History), 1971 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 3 December, 1971 Price 2 



LIZARDS AND SNAKES FROM TRANSJORDAN, 

RECENTLY ACQUIRED BY THE 
BRITISH MUSEUM (NATURAL HISTORY) 

By YEHUDAH L. WERNER 

CONTENTS 

Page 

SYNOPSIS ........... 215 

INTRODUCTION. .......... 215 

LOCALITIES ........... 216 

METHODS ........... 218 

LACERTILIA ........... 219 

Gekkonidae .......... 219 

Agamidae .......... 222 

Chamaeleonidae ......... 232 

Lacertidae .......... 234 

Scincidae .......... 241 

OPHIDIA ............ 243 

Colubridae . . . . . . . . . . 243 

DISCUSSION ........... 246 

General biogeography of Transjordan ..... 246 

Distribution of reptiles ........ 249 

ACKNOWLEDGEMENTS . . . . . . . . .251 

APPENDIX ........... 251 

REFERENCES ........... 254 

SYNOPSIS 

A report on 45 lizards and snakes, representing 23 species and subspecies, collected in northern 
and southwestern Transjordan, mostly during 1963-1965. Taxonomic characters are pre- 
sented, and compared with data from adjacent areas, mainly Cisjordan. Relevant Trans- 
jordanian specimens in the Hebrew University of Jerusalem are also considered, and some 
identifications are revised. Field observations are cited. Agama pallida haasi ssp. nov. is 
described (type: BM 1965.800; 18 paratypes in BM, HUJ, FMNH). The only additions, on 
the species level, to the Trans Jordanian fauna, are Coluber rhodorhachis Jan and Malpolon 
moilensis Reuss. The ecological and phytogeographical subdivision of Transjordan into 
Mediterranean, Irano-Turanian, and Saharo-Sindian territories is reviewed. The distribution of 
reptiles appears to accord with this subdivision. The difference between the herpetofaunas of 
Trans- and Cisjordan, on the specific and subspecific levels, is greater in the south than in the 
north. Notably 7 Irano-Turanian and Saharo-Sindian forms of Transjordan do not occur in 
Cisjordan. It is suggested that the Wadi 'Arava together with the steep mountains bordering 
it on the east, may constitute a barrier to the distribution of reptiles. 

INTRODUCTION 

TRANSJORDAN, or Eastern Palestine, is of great zoo-geographical interest. In the 
north-west it borders on the mesic (Mediterranean) regions of Cisjordan (or Western 
Palestine), and Syria. To the north and north-east its steppe is continuous with the 



216 Y. L. WERNER 

steppes of Syria and Iraq, while its south-eastern portions are part of the Arabian 
desert. In the south-west, along the Wadi 'Arava, Transjordan adjoins the arid 
south of Cisjordan (Negev of Isreal), which, through Sinai, affords communication 
with north-eastern Africa. 

Despite the efforts of numerous naturalists, zoologists and herpetologists, the 
herpetofauna of this whole region remains imperfectly known. The best-known 
territory is the part of Cisjordan which has been within Israel since 1948, although 
the latest review of its herpetofauna in a European language (Haas, 1951) is now 
outdated due to subsequent collecting. More recent information is available to 
readers of Hebrew (Barash and Hoofien, 1956; Wahrman, 1963; Y. L. Werner, 1966). 
The herpetofauna of Sinai was reviewed by Schmidt and Marx (1956) and Marx 
(1968), and that of Iraq by Khalaf (1959). The herpetofauna of Syria and Lebanon 
was the subject of several older reports (referred to by Flower, 1933; Schmidt, 1939; 
Haas, 1951), and one recent publication (Zinner, 1967). 

The least known territory, herpetologically, is Transjordan, where little collecting 
has been done. Apparently the only recent papers dealing specifically with the 
herpetofauna of this area are those of Schmidt (1930), Parker (1935), Haas (1943, 
1951), Hoofien (1965, 1969) and Werner (1968). A few reports of broader scope also 
deal with Transjordanian reptiles; notably those of Peracca (1894), Barbour (1914), 
Schmidt (1939) and Wettstein (1951), and those cited by these authors or by Parker 
(1935) and Haas (1943, 1951). 

Recently the British Museum (Natural History) obtained 39 specimens of lizards, 
and 6 of snakes, from Transjordan, thanks to the thoughtfulness of three parties 
whose primary object had not been the collection of preserved reptiles: Mr. S. 
Bisserot of the British Jordan Expeditions 1963 and 1965 (Mountfort, 1965), Mr. D. 
Western of the University of Leicester, and (one specimen) Mr. W. Larmuth. The 
present material makes a notable addition to our knowledge of the reptiles of Trans- 
Jordan and their distribution. 

In this paper all but one of .these new specimens* are described and discussed 
with a view to stimulating the interest of herpetologists in this little-known region. 
Comparisons are made with specimens previously collected in Transjordan by Prof. 
G. Haas, Prof. H. Mendelssohn and Mr. J. H. Hoofien (Haas, 1943, 1951 ; Hoofien, 
1957) and deposited at the Hebrew University of Jerusalem, and with series from 
Cisjordan in the same collection. The zoogeographical implications of the limited 
data available are discussed. 

LOCALITIES 

The localities are indicated by numbers in text-figures i and 2, as follows : 

1. Jordanian-Syrian border, 5. Wadi Ratam. 
Jerusalem-Damascus road. 6. El Azraq. 

2. Tell el Mukheizin. 7. Wadi Aseikhim. 

3. Ain el Enoquiya. 8. Azraq Shishan. 

4. Azraq Druz. 9. Jebel Uweinid. 

*A specimen of Stenodactylus grandiceps Haas, $, BM 1963.665, collected by S. Bisserot at the Azraq 
Oasis (in a sandy area), was received by the British Museum but not examined by the author. 



LIZARDS AND SNAKES FROM TRANSJORDAN 



217 



10. Qasr Amra. 

11. Tell Qarma. 

12. Qa el Umari. 

13. Shaubak. 

14. Wadi Musa. 



15. Petra. 

16. Basta. 

17. Rum. 

18. Aqaba. 



35 



-33 



-32 



-31 



-30; 



18 



36 



37 



38 



39 

\ _ 
\ 
\ 
\ 

\ 
\ 



AMMANo 



2* 



12. 



17 



50 



FIG. i. Localities from which material is reported here. The rectangle around 6 is en- 
larged in fig. 2. Locality names in text. (Political frontiers as before 1966.) 



218 



Y. L. WERNER 




FIG. 2. Localities near Azraq from which material is reported here. This area corresponds 
to the rectangle around 6 in fig. i. Locality names in text. B, Basalt areas; W, Water; 
MF, Mud flats. 



METHODS 

Abbreviations: 

BM British Museum (Natural History), London. 

FMNH Field Museum of Natural History, Chicago. 

HUJ-R Herpetological Collection, The Hebrew University of Jerusalem. 

SMF Senckenberg Museum, Frankfurt. 

SV Snout-vent length. 

%SV Percentage of snout-vent length. 

Measurements: SV length is from the tip of the snout to the anterior margin of 
the cloaca. Measurements of parts of the body are preferably expressed in %SV 
(see abbreviations, also Werner, 1969) rather than in absolute values. In Agama 
pallida, head length to occiput is from the tip of the snout to the palpated occipito- 
atlantal articulation, in a straight line (with dividers); and head length, total is 
from the tip of the snout to the rear of the angle of the jaws, parallel to the long 
axis of the animal. The head index is derived from the formula 100 x "Head 
length, total"/"Head width". 

In the case of Agama pallida, the statistical significance of differences in numerical 



LIZARDS AND SNAKES FROM TRANSJORDAN 219 

values between sample pairs was tested in the following manner: i. The signific- 
ance (a = 0-05) of the difference between the variances of the two samples was 
checked, using the F-distribution. 2. Regardless of the result, the statistic t 
(Winer, 1962 : 28) was computed, and used in Student's -test (Winer, 1962 : 28-29) 
as follows: 3. First, t was assigned n a -f n b 2 degrees of freedom; all cases in 
which sample means did not differ significantly, needed no further consideration. 
4. Where sample means differed significantly (a = 0-05), and the variance had not 
differed significantly, the result was accepted. 5. Where the means differed 
significantly, but the variances had also differed significantly, the significance of 
the difference between the means was confirmed by assigning to t only the degrees 
of freedom of the smaller of the two samples, making use of Welch's approximation 
(Winer, 1962 : 37-3$) to the i' distribution, (X a - X b ) - (H a - ft,)- 

V(s'/n a ) + (s2/n b ) 
Field, notes are credited expressly to the observer (collector) except where the 

observed animal is cited by its number, whereby the collector's name can be located 

in 'Material Examined'. 

Other terms and procedures are as explained by Haas and Werner (1969), else as 

defined by H. M. Smith (1946 : 17-30) or as presented by Peters (1964). 

LACERTILIA 

GEKKONIDAE 

Hemidactylus turcicus turcicus Linnaeus 

Lacerta turcica Linn6, 1758. Syst. Nat., ed. 10, i, p. 202 (Orient). 
Hemidactylus turcius, Boettger, 1876. Ber. offenbach. ver. Naturk., 15/16 : 57. 
Hemidactylus turcicus turcicus, Mertens, 1925. Abh. senckenberg. naturf. Ges., Frankfurt a.M., 
39 : 60. 

MATERIAL EXAMINED (i). BM 1965.683 $, [between Azraq Shishan and Azraq 
Druze],* April-May 1965, S. Bisserot. 

PHOLIDOSIS. Dorsal tubercles keeled, 14 in a diagonal row across the back. 
Usually 2-3 granules between successive or adjacent tubercles. Counts of scansors 
are presented in Table i. 

MEASUREMENTS. SV : 47 mm ; tail incomplete. 

COLORATION. Dorsum with five semi-regular longitudinal rows of brown mark- 
ings of irregular shape. 

FIELD NOTES. "The only other gecko found was the Turkish Gecko (Hemi- 
dactylus turcicus) [one of] which was found in a damaged condition on the track 
between Azraq Shishan and Azraq Druze" (S. Bisserot). 

REMARKS. Five specimens previously collected in Trans Jordan were available 
for comparison (HUJ-R 1063 70 km S. of Amman: Sisah-Chissa, 27 March 1936, 

*Not indicated on the specimen label, but derived from Mr. Bisserot's field notes. 



220 Y. L. WERNER 

Haas; HUJ-R 1064 Hissa-Ma'an, 28 March 1936, Haas; HUJ-R 1068 Wadi Daba'a 
SSE of Amman, July 1938, Haas; HUJ-R 1587 Jerash, 18 November 1945, Haas 
and Hoofien; HUJ-R 1588 Birketen- Jerash, 16 November 1945, Haas and Hoofien). 
Of these, four likewise have 14 rows of tubercles, but one has 12. Scansor counts 
for these specimens are included in Table i and compared to Boulenger's (1885) data 
for H. turcicus and H. sinaitus. The specimens from Transjordan, including those 
from the desert, are typical H. turcicus. 

TABLE i 

Scansor counts for Hemidactylus turcicus from Transjordan, compared to Boulenger's (1885) 
counts for H. turcicus and H. sinaitus. (N, number of specimens.) 

Locality (and repository of Scansors (single or paired) under :- 

material.) , * ^ 

Fingers Toes 







f 


- % 


t 


^ 




N 


First 


Fourth 


First 


Fourth 


Azraq area (BM) 


i 


7 


9 


7 


12 


Jerash area (HUJ) 


2 


6-7 


8 


6 


9-10 


Southern Transjordan (HUJ) 


3 


7-8 


8-9 


6-7 


IO-I2 



Transjordan, cumulative 6 6-8 8-9 6-7 9-12 

Boulenger's H. turcicus 28 6-8 8-10 6-8 9-11 

Boulenger's H . sinaitus 157 58 

A pattern of dark (brown) spots arranged in 6-8 longitudinal rows occurs in 
HUJ-R 1587-8 and, to a lesser extent, HUJ-R 1064. None show a pattern of 
crossbands or of large X-shaped designs (each stretched across the back), whereas 
these are often encountered in specimens from Cisjordan, the latter pattern being 
particularly common in certain localities (e.g. En Gedi). 

Ptyodactylus hasselquistii cf. guttatus von Heyden 

Ptyodactylus guttatus von Heyden, 1827, in Riippell, Atlas Reise nordl. Afrika, i. Zool.: Rep- 

tilien, p. 13, pi. 4, fig. i (Tor, Sinai). 
Ptyodactylus hasselquistii Phalanx gutlata (part), Anderson, 1898. Zoology of Egypt, i. 

Reptilia and Batrachia, London, pp. 65-67, pi. 6, figs 4, 5 and 5a, pi. 7, figs 6 and 7. 
Ptyodactylus hasselquistii guttatus, Barash and Hoofien, 1956. Reptiles of Israel, Tel-Aviv, 

p. 161. 

MATERIAL EXAMINED (4). BM 1963.664 $ Rum (rocky area), 1963, S. Bisserot. 
BM 1965.782 juv. Petra (on red sandstone), 2 August 1965, D. Western. BM 
1965.783 and 1965.784 juvs. Wadi Musa (under yellow sandstone rock), 4 August 
(3.30 pm and 12.30 pm resp.), D. Western. 



PHOLIDOSIS. Tubercles of the three juveniles flat-conical, with indications of 
keels ; of the adult, distinctly keeled. Hardly any tubercles in front of the ear or on 
the forearm. Other details in Table 2 ("Petra-Rum") which may be compared 
with Loveridge's table (1947 : 279). 



LIZARDS AND SNAKES FROM TRANSJORDAN 221 

MEASUREMENTS. Adult: SV: 68 mm; tail missing. Juveniles: SV: 55, 31 and 
30 mm resp. ; only the last with complete (?) tail: 23 mm. 

TABLE 2 

Pholidosis and measurements of 12 Ptyodactylus hasselquistii sspp. from Transjordan, for 
comparison with Loveridge's (1947 : 279) table. 

% 6 t % S 

DtfltuO 5 3 t"" 1 O I-" 

S -3 .S g d * 1 ^ 

'S 3 'd o fefe ^Sj3 

S J2 J9 g g & to 

^ O g .5 .5 o Is Is J 

Locality (and l| <u ^ ^ ^ g 

repository 2 c fc 1 | g 1 J 2 J S 

of the material) S fc - | | 1 I -a 

* * O, 30 * 38 -3 "3 

P i-3 H Q c/) W H H O c/5 G H 

Petra - Rum (BM) 4 i 3-4! 11-14 Ir -i3 2 12-13 3~4 9~u 4-6 68 ? 

S of Guveira (HUJ) 113 14 11-12 12 2-3 12 6 62-5 54 

Basalt Desert (BM) i i 3 12-13 IO 10 2 9 6 41 38-5 

Jerash area (HUJ) 6 i 3 10-12 8-10 12 2~5 3 g-n 4 6-7 67 52 

1 Four were observed on only one side of one specimen (BM 1965.784). 

2 But 15 were counted on one side of one specimen (BM 1963.644). 

3 Each of the extreme counts (2 and 5) occurred on only one side of one animal; in all other cases there 

were 3-4 scansors under the ist toe. 

4 The count is uncertain on one side of HUJ-R 1653 (? 7-9). 

COLORATION. Collector's notes: BM 1965.782, "grey with green and brown 
spots, light orange mottling". BM 1965.783 and 1965.784, "beige with brown and 
white spots". After preservation these three juveniles from near Petra are a light 
greyish brown, with small light spots which are nearly round, and are 3-6 granules 
across. They are arranged fairly regularly in longitudinal rows ; dark brown spots 
of less regular shape are arranged between them. The adult from Rum (BM 
1963.664) is coloured similarly, but the light spots are only faintly discernible, and 
the dark spots are larger, of more irregular shape, and less regularly scattered. All 
specimens have whitish underparts. 

REMARKS. Ptyodactylus hasselquistii is notorious for its high geographical vari- 
ability (Flower, 1933; Loveridge, 1947; Werner, 1965). This is particularly true in 
the regions surrounding the Gulf of Suez, Gulf of 'Aqaba, and the Dead Sea, where 
the typical form meets, mixes, or intergrades with guttatus. The present series 
shows points of resemblance to guttatus from (central) Israel, and to the original 
illustration of von Heyden ; but on the basis of our single adult without tail, alloca- 
tion remains uncertain. 

One subadult from S of Guveira (HUJ-R 1027, 28 March 1936, Haas) resembles the 
present specimens in its keeled dorsal tubercles and in lacking tubercles in front of 
the ear and on the forearm. Its particulars are included in Table 2. 



222 Y. L. WERNER 

Ptyodactylus hasselquistii puiseuxi Boutan 

Ptyodactylus puiseuxi Boutan, 1893. Rev. Biol. du Nord de la France, 5 (9) : 27-32, pi. 3, fig. 4 

("Bords du lac de Houleh"). 

Ptyodactylus lobatus syriacus, Peracca 1894. Boll. Mus. Zool. Anat. comp. Torino, 9 (167) : 

1-6 (Jerash, Transjordan). 

Ptyodactylus hasselquistii puisieuxi, Haas, 1951. Bull. Research Counc. of Israel, i (3) : 95. 
Ptyodactylus hasselquistii puiseuxi, Barash and Hoofien, 1956. Reptiles of Israel, pp. 160-161. 

MATERIAL EXAMINED (i). BM 1965.682 juv. Basalt desert, Wadi Aseikhim, 
April-May 1965, S. Bisserot. 

PHOLIDOSIS AND MEASUREMENTS. Tubercles not keeled, each resembling a low 
cone. Between ear and corner of mouth 10-15 tubercles, and on each forearm 
about 15. Other details in Table 2 ("basalt desert") which may be compared with 
Loveridge's table (1947 : 279). 

COLORATION. After preservation, dark brownish grey with round whitish dots 
(2-3 granules in diameter) alternating with roundish dark spots (5-8 granules in 
diameter). Underparts light grey. Tail with conspicuous alternating dark and 
light half rings on the dorsal surface ; ventral surface is grey with whitish mottling. 

FIELD NOTES. ' . . . the fan-footed gecko (Ptyodactylus hasselquistii) . . . 
greater quantities were found when they eventually were observed for the first 
time on April 3oth 1965, always on the basalt. One clutch of eleven eggs [obviously 
at least 5| clutches] of this species was found at Wadi Aseikhim, nine of which were 
hatched and two not. One captive specimen laid two eggs during the journey 
back . . . " (S. Bisserot). 

REMARKS. The same subspecies occurs at and around Jerash (terra typica, 
Peracca, 1894) as also shown by 6 specimens collected by Haas and Hoofien in 1945 
(HUJ-R 1651-5 and 6112). Particulars of these are included in Table 2. The 
present specimen, closely resembling puiseuxi from Jerash and from northernmost 
Cisjordan, allows us to suggest' that puiseuxi is probably primarily associated with 
basalt rocks, regardless of whether these are in a mesic habitat (northern Cisjordan 
and northwestern Transjordan) or in an arid one (basalt desert of northeastern 
Transjordan). It is not, however, absolutely restricted to basalt, occurring also on 
adjacent calcareous formations. 

AGAMIDAE 

Agama pallida haasi subsp. nov. 
(Text-fig. 3; Pis i, 2) 

Agama ruderata pallida (part), Haas, 1943. Copeia i : 12. 

Agama pallida (part), Haas, 1951. Bull. Res. Counc. Israel i (3) : 72-74. 

HOLOTYPE. BM 1965.800 $ Azraq in Transjordan, 12 August 1965, D. Western. 

PARATYPES (18). Males (10): BM 1965.684 Jebel Uweinid (Basalt desert), April- 
May 1965, S. Bisserot; BM 1965.796 Azraq, 12 August 1965, D. Western; HUJ-R 
1117 between Sisah and Ma'an, March 1936, G. Haas; HUJ-R 1121 N. Dahaa, 65 m 



LIZARDS AND SNAKES FROM TRANSJORDAN 223 

SSE Amman, June-July 1938, collector unknown; HUJ-R 1134 between Hissa and 
Amman, 28 March 1936, G. Haas; HUJ-R 5215, 5216 and 5217 between Sisah and 
Ma'an, March 1936, G. Haas; HUJ-R 1227 near Palmyra, Syria, June 1944, Theodor; 
FMNH 48468 Wadi Dabaa 65 m SSE Amman, July 1938, collector unknown (from 
Hebrew University). Females (4): HUJ-R 1118 between Hissa and Ma'an, nr. 
Ma'an, 28 March 1936, G. Haas; HUJ-R 1120 60 km NE Zerka, Transjordan (no 
date), Sjoma Graber; HUJ-R 1124 about 12 km S of Amman, 26 March 1963 (?) 
Mendelssohn; HUJ-R 1884, Wadi Debba, Transjordan, Summer 1939, collector 
unknown. Juveniles (4): BM 1936.666 Azraq, 16 April 1963, S. Bisserot; BM 
1965.797, 1965.798 and 1965.799 Azraq, 12-13 August 1965, D. Western. 

DIAGNOSIS. Ear opening distinctly longer than high, bordered above by a row 
of conspicuous spines; not round with fairly smooth margin as in Agama pallida 
pallida Reuss 1833 from eastern Egypt, Sinai, and southern Israel. Total size 
larger, head and body more elongate than in A. p. pallida. Ventral scales usually 
smooth, not keeled as in Agama agnetae F. Werner 1939 from western Iraq. 
(Pis i & 2). 

DESCRIPTION OF HOLOTYPE. A male. Head very convex, short and thick, but 
distinctly longer than broad (head index: 114). Nostril not tubular, superior, barely 
above the indistinct canthus rostralis. Nasal shield flat. Upper head scales convex, 
with short terminal keels ; occipital not enlarged. No well-developed spines on the 
hinder part of the head, but a few occipital scales are pointed (resembling the 
enlarged scales scattered on the back). A fringe of 3-4 distinct spines on the upper 
edge of the ear, pointing downwards (in the preserved specimen), except for one 
spine, on the anterior margin of one ear, which points backwards. Ear opening 
smaller than eye opening, elongate, nearly twice as long as high; its upper (spiny) 
border nearly straight and horizontal (PI. i B). No gular pouch. Body depressed, 
not as short as in A. pallida pallida (Pis i A; 2 A). Dorsal scales very small, 
irregular, faintly imbricate, indistinctly keeled; intermixed with scattered larger 
scales each of which bears a short keel, sometimes ending in a short spine. Scalation 
of limbs, and proximal quarter df tail, similar to that of back, but the small ground 
scales larger than on back. Ventral scales smooth, imbricate. Tibia longer than 
the skull (to occiput). Third ringer shorter than fourth, fifth not extending as far as 
second; third toe much shorter than fourth, fifth not extending as far as first. Tail 
I 45% SV long (somewhat more than twice as long as the distance from gular fold 

MEASUREMENTS OF HOLOTYPE. mm 

Total length 189 

Snout-vent 77 

Head length (to occiput) 20 

Head length (total) 24 

Width of head 21 

Body (occiput-vent) 57 

Forelimb 40 

Hindlimb 59 

Tail 112 



224 Y - L. WERNER 

to vent), circular in cross section, its distal three quarters with subequal keeled 
scales. A double row of 'anal pores' (10 -f 12). 

COLORATION OF HOLOTYPE. Collector's note: "mottled dark grey brown: white 
dashes". After preservation, brownish grey. Pileus yellowish. In the orbital 
area, below the eye, six faint grey radiating streaks; side of head otherwise plain. 
Dorsum with four darker brown crossbands, each interrupted by an irregular whitish 
vertebral streak. First crossband in front of, second behind, shoulder. Third, 
indistinct. Fourth just in front of pelvis. Thirteen uninterrupted crossbands on 
tail. The first two are similar in colour to the ones on the body, the remainder are 
paler. Underparts light cream, throat mottled with 8-10 grey wavy longitudinal 
bands. 

VARIATION OF THE TYPE SERIES. Females have no anal pores, and their heads 
are shorter (relative to SV length) than in males. The largest specimen is a female 
(HUJ-R 1118), SV 93-5 mm; largest male (HUJ-R 1117) SV 89 mm. At the upper 
border of the ear there are 2-4 large and 0-2 small spines ; the commonest arrange- 
ment is 3 large and i small spine. All juveniles, including the smallest (SV: 33 mm), 
show the distinctive ear features of the new form, except that the enlarged scales 
bordering the ear opening dorsally are not spiny (PI. i C-D). The ventral scales 
are moderately keeled on the posterior abdomen of one specimen (HUJ-R 1134). 
The variations of pholidosis, measurements and proportions, and pattern, and com- 
parable variations of A. p. pallida from southern Cisjordan are summarized in Table 
3. Differences between the samples from Trans Jordan and Cisjordan (comparisons 
being made separately among males, females and juveniles), were statistically 
significant only in the following instances: Among males, the two samples differed 
significantly in SV length (t = 6-32; tio (-05) = 2-23) ; in head length (to occiput) in 
%SV (t = 2-59; t 9 (-05) = 2-26); in head width in %SV (t = 4-53; t 28 (-05) = 2-05); 
and in the head index (t = 3-52; t28 (-05) = 2-05). Among females the two samples 
differed only in head width in %SV (t = 3-18; tzi (-05) 2-08) and in the head 
index (t = 2-13; t 2 i ( . 5) = 2-08). 

Collector's note on coloration: "The Pale Agamid (Agama pallida) varied con- 
siderably, in colour and markings .... A. pallida showed no colour changes 
under any circumstances". (S. Bisserot.) 

FIELD NOTES. "Three species of ... Agamidae were seen and collected .... 
A. pallida was the most common but was found only on the hamada. . . . On 
the hamada areas the dominant reptiles appeared to be ... and Agama pall- 
ida ... " (S. Bisserot). Two of the adult males (BM 1965.800 and 1965.796) 
were caught "among small rough stones: flint, basalt and chert on brown silty 
matrix between stones" (the first at 11.30 h). A juvenile, BM 1965.798 "in shade 
under Holoxocum silicanum [? Haloxylon salicornicum] near black basalt rocks 
(12.30 pm)"; another juvenile, BM 1965.797, "on flint stone desert", and another 
among "basalt outcrops large basalt boulders with white interdispersed silt under 
rock". HUJ-R 1134 was caught on "Ebene Stein Wiiste" (even stone desert). 

"The Pale Agamid . . . appeared to rely on three methods of escape, firstly by 
quick bursts of running when the body was held high off the ground and the head up, 



LIZARDS AND SNAKES FROM TRANSJORDAN 



225 



than by flattening the body to the ground and remaining motionless, relying on 
camouflage, and lastly by an aggressive stance with mouth open always facing the 
attacker." (S. Bisserot.) 

GROWTH. The three juveniles collected on 12-13 August 1965 by Mr. Western 
measure 44, 38-5 and 33 mm respectively (SV). These obviously had hatched earlier 
in the same season. The one taken on 16 April 1963 by Mr. Bisserot measures (SV) 
46-5 mm and evidently had hatched in the previous summer. The 17 specimens for 
which the date (at least the month) of collection is known (Text-fig. 3) make it 
probable that in Transjordan the hatchlings of the year reach ca. 40-50 mm (SV) by 
autumn, grow to ca. 70-80 mm during the following year, and attain 80-90 mm in 
their third warm season. 



90- 



Z 70- 

LLJ 
> 

I 

O 60- 

z 

00 

50- 



40- 



30- 



a 
o 



O 
O 

o 



j 'F'M'A'M'JJ'A'S'O'N'D' 

DATE OF COLLECTION, MONTHS 

FIG. 3. Agama pallida haasi subsp. nov. Sizes of animals caught at different times of 
the year. Squares, males; Circles, females; Diamonds, juveniles. Open symbols, 
animals from Transjordan; Solid symbol, animal from near Palmyra, Syria. 

REMARKS. It is a pleasure to name this lizard in honour of Professor Georg Haas 
who had already commented on its relatively large size (1951). The new taxon is 
most closely allied to Agama pallida Reuss 1833 from Eastern Egypt, Sinai and 
Southern Cisjordan. In Reuss' original description there is no indication of the 
type locality, except that the whole material under discussion had been collected by 



226 



Y. L. WERNER 



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LIZARDS AND SNAKES FROM TRANSJORDAN 



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228 Y. L. WERNER 

Riippell. Reuss' original specimen label indicates "Aegypt, super." (Klemmer, 
1967). The species, however, does not seem to occur in Upper Egypt, and Anderson 
(1896 : 79) gives "Sinai" as the type locality (see also Flower, 1933). Through the 
courtesy of Dr. Klemmer I could examine excellent photographs of the type (SMF 
10007) an d satisfy myself that material from southern Cis Jordan and eastern Egypt 
(Kassassin) is in good general agreement with it. Reuss' description includes no 
scale counts, but some of his many measurements are represented in Table 3. 

The new form is so far known from central and northern Transjordan and SE 
Syria, but specimens from the rest of Syria and from Iraq will probably also be found 
to belong to this form. A. p. pallida Reuss and A. p. haasi n. ssp. are allopatric, 
possibly separated by the steep slopes constituting the eastern rim of the Wadi 
'Arava. Though I have seen no intermediate forms, the few specimens from the 
Wadi 'Arava being typical pallida, conclusive evidence of reproductive isolation is 
not yet available. Thus it seems best to accord the two forms subspecific rank. 

In the northern part of its range A. p. haasi may be sympatric with A. ruder ata 
Olivier. I have omitted the references for one of these forms having been recorded 
within the accepted range of the other, as probably some of these instances are based 
on mis-identifications. However, it is my impression that this is not so in all cases 
(see also Pasteur and Bons, 1960). It is interesting that it is easier to distinguish 
between A. p. haasi and A. ruderata, on the basis of general habitus, than between 
A. p. pallida and A. ruderata. Moreover, sexual dimorphism in SV length is 
apparently moderate in A. p. haasi (largest $, 89 mm (HUJ-R 1117); largest $, 
93-5 mm (HUJ-R 1118)), whereas it is considerable in both A. ruderata (Pasteur and 
Bons, 1960) and A. p. pallida (among 117 specimens from Cisjordan in the Hebrew 
University collection, largest $, 75 mm (HUJ-R 7509) ; largest $, 87 mm (HUJ-R 
5506)). These phenomena apparently represent a case of "sympatric character 
divergence" (Mayr, 1965 : 82). 

It has been argued that A. pallida Reuss 1833 is conspecific with, and indistin- 
guishable from, A. mutabilis Merrem 1820 (Pasteur and Bons, 1960; Wermuth, 1967). 
In fact, this possibility had already been mentioned by Anderson (1898). However, 
the arguments (and diagrams) of Pasteur and Bons do not entirely exclude the 
possibilities that these are either two distinguishable allopatric forms (Flower, 1933) 
(with a complex borderline, or intergrading), or even sibling (partly sympatric) 
species (Schmidt and Marx, 1956 : 25). It therefore seems most prudent to retain, 
at present, the specific name pallida for the populations to which it has traditionally 
been applied. 



Agama blanfordi fieldi Haas and Werner 

(PI. 3 A) 

Agama persica fieldi Haas and Werner, 1969. Bull. Mus. Comp. Zool. Harvard, 138 (6) : 
337-339, pis. 2-6. (Saudi Arabia: Al-Caissumah - Turaif .) 

MATERIAL EXAMINED (i). BM 1965.686 $ Qa el Umari (hard sand desert), 1965, 
S. Bisserot. 



LIZARDS AND SNAKES FROM TRANSJORDAN 229 

PHOLIDOSIS. Dorsal scales subequal, keeled and shortly mucronate. Lateral 
scales similar but smaller. Ventral scales feebly keeled. Scales around middle of 
body, 80. 

MEASUREMENTS. SV, 105 mm; tail, 163 mm. 

COLORATION. Collector's note: "A. persica when first captured turned to a 
brilliant blue in the area of the dew-lap under the chin but not on any other part of 
the body. Both A. sinaita and A. persica turned blue when killed and preserved in 
spirit." (S. Bisserot.) Yellowish grey*. Dorsally, four darker longitudinal bands, 
brown with still darker margins. Each band is of uneven width, and contains about 
six alternating dilated and constricted zones. Through the dilated zones run trans- 
verse series of white dots which are interrupted by a median light band. Each dot 
coincides with an enlarged scale, making it more conspicuous. On the head there 
are two brown wavy crossbands, preceded by a longitudinal patch of the same colour. 
On the tail the dorsal pattern gradually changes to one of simple dark rings. Belly 
with a distinct central longitudinal, grey band, and irregular lateral ones. Gular 
pouch dark grey with remnants of blue. 

FIELD NOTES. "Three species of ... Agamidae were seen and collected . . . 
on the flat dried sand areas only the one specimen of A. persica was found". S. 
Bisserot. 

REMARKS. This specimen is identifiable according to Boulenger's (1885) key as 
A. blanfordi (S. C. Anderson, I966a; nom. subs, for A. persica Blanford 1881, nom. 
preoccup.) because of its unequal dorsal scales. However, the gular pouch and the 
scattered enlarged scales are less developed than in blanfordi and the head and body, 
especially the former, are more depressed than in this form. On the other hand, a 
series of very similar specimens collected in NE Saudi Arabia by Mr. Henry Field 
shows considerable variation in the development of the enlarged dorsal tubercles; 
several specimens have homogenous scaling and are thus identifiable as isolepis 
(Boulenger, 1885, lectotype from between Magas and Bampur, southeastern Iran 
S. C. Anderson, i966b). Similar, apparently, were the two specimens from Meso- 
potamia which Steindachner (1917) identified as "isolepis with a unique pattern". 
Our specimen is very similar to the pair depicted by him (at least in proportions, 
pattern, and non-meristic scale characters). 

Apparently this is a form allied both to the agilis-isolepis Rassenkreis (Wettstein, 
1951) and to blanfordi. It is, in certain respects, intermediate between the two. It 
is characterized by variably (mostly feebly) developed dorsal tubercles and gular 
pouch, and by a very distinctive pattern of longitudinal bands. A confusion con- 
cerning agilis and blanfordi had already been suspected by Schmidt (1941). 

This new form has previously been referred to as A. persica Blanford. Thus at 
least part of the series mentioned by Haas (1957) belongs to A. blanfordi fieldi, as 
judged by specimen CAS 84541 (now HUJ-R 7081) and by his description of the 
pattern of CAS 84477. The latter description has been accepted by Khalaf (1959) 

*When examined by the author. 



230 Y. L. WERNER 

as applying to A. persica Blanford though differing from that usually encountered 
in this species. 

Agama sinaita von Heyden 

(PI. 3 B-E) 
Agama sinaita von Heyden, 1827. In Riippell, Atlas Reise nord. Afrika, Kept., p. 10, pi. 3 (Sinai). 

MATERIAL EXAMINED (4). BM 1965.685 <$ Wadi Ratan [W. Ratam] (basalt 
desert), April-May 1965, S. Bisserot. BM 1965.801 <f>, BM 1965.802 $ Petra (on red 
and yellow sandstone respectively), 2 August 1965, D. Western. BM 1965.803 $ 
Wadi Musa, near Petra (basking on soil on top of rock), 4 August 1965, D Western. 

PHOLIDOSIS. In the specimen from the basalt desert (BM 1965.685) the tail is 
moderately compressed laterally including its thick basal portion. The two dorsal 
rows of caudal scales are enlarged, their thick keels creating the impression of a 
slight crest. The specimens from Petra and Wadi Musa resemble specimens from 
Cisjordan in that the thick part of the tail is nearly cylindrical, and carries dorsally 
four straight rows of strongly keeled scales, the two median rows being little better 
developed than their immediate neighbours (PI. 3 C & E). 

MEASUREMENTS. SV: 103, 69, 79, 79 mm. Tail of the last: 125 mm (other tails 
incomplete). In all specimens the third toe is hardly longer than the fourth (PI. 3 B). 

COLORATION. "The ability of A. sinaita to change colour was observed on several 
occasions but did not appear to follow a regular pattern. Most specimens were 
observed to be a brilliant cobalt blue on first sight [Mountfort, 1965 : plate 4oa] but 
changed to a dark chocolate brown when pursued. One specimen kept alive changed 
from brown to blue over the head and shoulders and part of the flanks when food was 
put in its mouth. This reaction was repeated in captivity in this country [England] 
and was also caused by the temperature being raised to 80 F [27 C] or higher . . . 
A. sinaita turned blue when killed and preserved in spirit." (S. Bisserot.) 

Professor H. Mendelssohn (Tel- Aviv University) has studied the colour changes of 
this species, as part of his research of its behaviour, and found marked sexual 
dichromatism. Hence the arrangement of the following notes on our specimens. 

Males: BM 1965.865 (preserved) : blueish grey (throat and belly darker), tail grey 
(yellowish ventrally). 1965.802, collector's note: "bright blue all over, faded after 
death." The preserved specimen is dark grey, nearly black, with underparts lighter 
(and posteriorly very light) brownish grey. 

Females'. Collector's notes summarized: "head blue when alive, turning brighter 
blue when killed. Body grey with orange blotches". After preservation the heads 
are blackish, the bodies dark grey, and the tails have alternating darker and lighter 
transverse bands. The throats are grey (reticulated in 1965.801) and the abdomens 
steel grey; the remaining underparts are cream- white. 

REMARKS. The unusually large male with compressed tail, from the basalt 
desert, conceivably represents a distinct subspecies, but this cannot be assessed on 
the basis of a single specimen. (Wettstein, 1951 : 433, mentions three specimens 



LIZARDS AND SNAKES FROM TRANSJORDAN 231 

from northern Trans Jordan, but gives no particulars apart from the colour.) The 
remaining specimens have tails similar to those of animals from southeastern Israel 
and northeastern Sinai, and the same is true of two collected by Haas between 
Guweira and Aqaba in 1936 (HUJ-R 1137, HUJ-R 5231). On the other hand, all 
the specimens from Trans] ordan are characterized by the third toe being hardly 
longer than the fourth. Specimens from southeastern Israel and northeastern Sinai 
usually have a longer 3rd toe (PI. 3 B & D). 

Agama stellio brachydactyla Haas 
(Text-figs 4, 5) 

Agama stellio brachydactyla Haas, 1951. Ann. Mag. Nat. Hist. (Ser. 12), 4 : 1052 (Israel : foot 
of Jebel Lussan, near Israel-Sinai frontier, S.S.W. of Beer-Sheba). 

MATERIAL EXAMINED (3). BM 1965.787-789, Basta (remarks on habitats, under 
coloration), 3 August 1965, D. Western. 

PHOLIDOSIS. A mid-dorsal band of unequal enlarged scales, about six times as 
broad as one of the larger scales. The transverse series of tubercles extend across 
this band. They are slightly interrupted medially but here some of the interstitial 
scales are almost as large as the tubercles themselves. All large scales are either 
distinctly keeled, mucronate, or spinous (Text-fig. 4). 




FIG. 4. Agama stellio brachydactyla. Mid-dorsal sclaes of BM 1965.787 from southern 
Transjordan (Basta). (From a photograph.) Scale, 10 mm. 

Lamellae under ist finger: 9, 10, 9. Under 4th toe: 22, 24, 22. 

MEASUREMENTS. SV: no, 84, 103 mm. Tail of smallest: 116 mm; other tails 
incomplete. Foot hardly longer than tibia, being shortest in largest specimen. 

COLORATION. Collectors notes: 1965.787, "Back: black; black and yellow trans- 
versely striped tail; orange blotches near head (basking on orange mauve quarzite, 



232 Y. L. WERNER 

3.25 pm)." 1965.788, "Yellow and black with orange spots on neck. (On yellow 
sandstone, 4.20 pm)". 1965.789, "Brownish with orange and brown blotches. 
Tail black with orange transverse stripes. (On soil besides flintstone, 3 pm)". 

In the preserved condition all three specimens are grey dorsally with pale yellowish 
blotches, the largest blotches arranged in mid-dorsal asymmetrical pairs, each pair 
tending to fuse and to form a large obliquely transverse blotch. On the tail, trans- 
verse bands of the same yellowish colour, which also covers the underparts, the 
throat being faintly reticulated with pale grey. 

REMARKS. All three specimens are presumably not fully grown. These indi- 
viduals are not very typical brachydadyla. In the number of lamellae under the 
toes, as well as in the relative size of the mid-dorsal scales, they rather resemble 
specimens from the northern Negev in Israel, which are intermediate between 
brachydadyla and the form inhabiting mediterranean Israel. The specimens from 
Basta are assigned here to brachydadyla in accordance with their coloration and also 
in order to indicate their geographical affinities (Daan, 1967). 

Six other specimens from Transjordan are available. Only one from Petra 
(HUJ-R 1096, 29 March 1935, Haas) shows a similar arrangement of a mid-dorsal 
band of subequal enlarged scales. Two other specimens from southern Transjordan 
(HUJ-R 1094 and 1103) and three from the Jerash-Amman area (HUJ-R noi, 
1 1 10 A, and mo B) have the dorsal transverse series of tubercles clearly separated 
by smaller scales, as is usual in specimens from northern Cisjordan. 

The number of lamellae under the 4th toe in the nine specimens from the two areas 
in Transjordan is presented in Text-fig. 5 which also includes, for comparison, 
samples from five localities in Cisjordan. As the figure shows, in Cisjordan there is 
a pronounced north-south gradient in this character (with the higher values in the 
north). A parallel but less prominent gradient is indicated in Transjordan. 



CHAMAELEONIDAE 

Chamaeleo chamaeleon recticrista Boettger 

Lacerta chamaeleon Linnaeus, 1758. Syst. Nat., p. 204 (Africa and Asia.) 

Chamaelo vulgaris var. recticrista Boettger, 1880. Jahresber. senckenberg. naturf. Ges. 
Frankfurt, p. 198 (Jeruslaem and Haifa). 

MATERIAL EXAMINED (i). BM 1963.667 juv. between Shaubak and Tafila (on 
camel thorn), 1963, S. Bisserdt. 



FIG 5. Agama stellio subspp. Numbers of lamellae under fourth toe in Trans- and Cisjor- 
dan. Material included in each sample was collected within the area of the map covered by 
the relevant circle. (A) Jerash-Amman area; (B) Desert locality included in sample C; 
(C) Petra-Basta area; (D) Hills surrounding Lake Tiberias; (E) Ramot-HaShavim ; 
(F) Jerusalem and adjacent Judaean Hills; (G) Be'er-Sheva and vicinity; (H) Sde- 
Boker-'Avdat area; M, Mean; N, Number of specimens. (Political frontiers as before 
1966.) 



LIZARDS AND SNAKES FROM TRANSJORDAN 



233 




234 Y - L. WERNER 

PHOLIDOSIS. Gular and abdominal crest of enlarged scales present. 
MEASUREMENTS. SV: 44 mm; tail: 42 mm. 

COLORATION. Grey with irregular dots of darker grey. On each flank two longi- 
tudinal rows of five light cream blotches. Enlarged scales of dorsal, gular and 
abdominal crests the same light cream. 

REMARKS. A half -grown specimen from El-Hamma (NW Transjordan, in Israel 
HUJ-R 1501; 13 March 1945, Coll. G. Haas) is similarly coloured. In a juvenile 
from Jerash (HUJ-R 1502; 15 November 1945, Coll. Haas and Hoofien), measuring 
36 mm (SV) the occipital casque is not (yet) developed. All three specimens conform 
to the chamaeleons of northern Israel rather than to the C. c. musae-like animals 
from further south (Hoofien, 1964). 

LACERTIDAE 

Acanthodactylus boskianus asper Audouin 

Lacerta aspera Audouin, 1829. Descr. Egypte, Kept., Suppl., p. 173, pi. i fig. 9 (Egypt). 
Acanthodactylus boskianus var. asper Lataste, 1885. Ann. Mus. Geneva 2 (2) : 496. 

MATERIAL EXAMINED (3). BM 1965.691 $ and BM 1965.693 $ Tell Quarma 
[=Tell Qarma] (blown sand wadi), April-May 1965, S. Bisserot. BM 1965.804 $ 
Wadi Musa near Petra (sandy soil near bushes), 4 August 1965, D. Western. 

PHOLIDOSIS. Scales across middle of body: 31, 38, but 55 in BM 1965.693. 
Gular scales in straight median series: 26, 32, 26. Lamellae under 4th toe: 21-22. 

MEASUREMENTS. BM 1965.691, 56 mm SV, 121 mm tail. BM 1965.693, 81 mm 
SV, and BM 1965.804, 62 mm SV (tails incomplete). 

COLORATION. BM 1965.804, collectors note: "brown: long orange stripes, 
spotted with black". 

REMARKS. Eleven other specimens from various localities in southern Trans- 
Jordan (HUJ-R 1335, 1338, 1341, 1661, 5041-6, 5053) have the following pholidotic 
counts: Scales across middle of back, 29-51 (against 29-42 in southern Cisjordan, 
N = 24). Gular scales in a straight median line, 25-35 (24-31, in Cisjordan, 
N =24). Lamellae under 4th toe, 19-22 (as in Cisjordan). Other conventional 
counts are also similar in Cis- and Transjordan except that some Cisjordan specimens 
show a reduction of the lateralmost ventral plates, so that only 8-9 longitudinal 
series are present. 

In general, adult male A. b. asper are larger than females. The 2 largest males 
seen from Transjordan are 75 and 77 mm (SV), so that the female from Tel-Qarma 
(81 mm) appears unusually large. 

The dorsal pattern of some Transjordan males includes rows of sharply defined 
blackish dots, instead of the more usual rows of irregular brownish spots. 



LIZARDS AND SNAKES FROM TRANSJORDAN 235 

Acanthodactylus grandis Boulenger 
(PI. 4 A, B) 

Acanthodactylus grandis Boulenger, 1909. Ann. Mag. Nat. Hist., 4 (8) : 189 (Jerud and Ataiba, 
Syria) . 

MATERIAL EXAMINED (2). BM 1965.692 $ Ain el-Enoquiyya (sand and stone 
wadi) ; BM 1965.694 $ Tell el Mukheizin (Hamada, beneath carcass of dog) ; both 
April-May 1965, S. Bisserot. 

PHOLIDOSIS. Scales across middle of body: 60; 60. In BM 1965.692 there is a 
fifth small upper labial before the center of the eye, on each side. 

MEASUREMENTS. BM 1965.692: SV: 73 mm; tail: 126 mm. BM 1965.694: 
SV: 96 mm (tail incomplete). 

COLORATION. BM 1965.692 (PI. 4 B) : Black spots, each covering up to ten scales, 
arranged in ten regular longitudinal rows, and in irregular transverse series. The 
two median rows begin at the occiput but disappear before the middle of the back ; 
4th and 5th row on each side present on flanks but absent from neck. Some rows 
extend on the tail, on the lateral sides of which the spots are represented as vertical 
blotches at every second suture between scale rings. Ground colour (preserved), 
nearly uniform grey (compare PI. VI of Boulenger, 1923). 

BM 1965.694 (PI. 4 A) : Six dark longitudinal stripes faintly indicated on back. 
Along these there are a few, irregularly scattered, small blackish spots, each covering 
up to 7 scales. 

In both specimens the sides of the head bear alternate light and dark vertical bars, 
one of the latter passes through the eye. 

REMARKS. All characters of both specimens are within the range of variation 
shown by A . grandis in the HU J collection, some of which have been mentioned by 
Haas (1943; Transjordan between Hissa and Ma'an). There is some difficulty in 
distinguishing immature A. grandis from A. scutellatus scutellatus, which likewise 
has smooth scales, since the range of variation of almost all conventional scale 
counts is nearly identical (see also Boulenger, 1923 : 50). BM 1965.692 resembles 
A. s. scutellatus in the number of supralabials (5) before the center of the eye, and in 
its relatively long foot with moderately well developed pectination. The specimen 
however is certainly assignable to A . grandis for the following reasons : 

The snout with its somewhat swollen nasals resembles that of other A . grandis, and 
not at all the pointed snout of A . s. scutellatus. 

While five supralabials in front of the center of the eye are characteristic of A. s. 
scutellatus, and 4 of A. grandis, 5 may sometimes occur in the latter (see also 
Boulenger, 1921 : 114-115). 

The longer foot and relatively stronger pectination (when compared to large A. 
grandis such as BM 1965.694) appear to be largely juvenile characters, which are 
paralleled in series of other species of Acanthodactylus containing mature and 
immature specimens. Moreover the pectination is still far less developed than in 
adult A. s. scutellatus from southern Israel. 



236 Y. L. WERNER 

The pattern conforms closely to that of A. grandis and differs most strikingly 
from that of A . s. scutellatus, as the latter never show any longitudinal arrangement 
of the markings. 

Acanthodactylus tristrami tristrami Gunther 
(PI. 4 C, D) 

Zootoca tristrami Gunther, 1864. Proc. Zool. Soc., p. 491 (Lebanon). 
Acanthodactylus tristrami Boulenger, 1881. Proc. Zool. Soc., p. 746, pi. 64, fig. i. 

MATERIAL EXAMINED (i). BM 1962.352 <j> Jordanian-Syrian border, Jerusalem- 
Damascus Rd (Outside customs shed) [probably loc. i on map], 4 June 1952, W. 
Larmuth. 

PHOLIDOSIS. Scales across middle of body: 59. Longitudinal rows of ventrals: 
ii. Other characters also in agreement with Boulenger's (1921) data for A. 
tristrami. 

MEASUREMENTS. SV: 82 mm; tail (tip missing) : 86 mm. 

COLORATION. The blackish markings tend to form a reticulum along each side 
of the dorsum (PI. 4 C). 

FIELD NOTES. "Died while ovipositing". 

REMARKS. Angel (1936) described from NE Syria A. t. orientalis (48-56 scales 
across middle of body), which was also reported from the neighbourhoods of Rutba 
(Schmidt, 1939) and Mosul (Haas, 1952) in Iraq. From Haditha, Iraq, Schmidt 
(1939) described A. t. iracensis (45-46 scales across middle of body). Thus 
Giinther's (1864) and Boulenger's (1921) A. tristrami was accorded subspecific rank 
as A. t. tristrami (58-65 scales across middle of body). 

Our specimen appears assignable to the typical form, as well as two specimens in 
the HUJ collection, reported by Haas (1943) : HUJ-R 1333 <$, 15 km S of Amman, 
SV 92 mm; HUJ-R 1332 <j>, 45 km S of Amman, SV 69 mm (after a year in captivity). 
Scales across middle of body, 58, 57 respectively. Ventrals in 10 rows. 

All three specimens are larger than Angel's A. t. orientalis (1936; 50-66 mm snout- 
vent, N = 8), although this alone would not have been taxonomically significant. 

The dorsal pattern of HUJ-R 1333 (<) consists of distinctly X-shaped blackish 
marks (PI. 4 D; Boulenger, 1921). Markings intermediate between this pattern and 
the reticulum of BM 1962.352 (?) are present on the female (the type ?) figured by 
Tristram (1885: PI. 16, fig. 2). 

Eremias brevirostris microlepis Angel 

Eremias brevirostris microlepis Angel, 1936. Bull. Inst. Egypte 38 : 112-113 ("Haouarine" 
5 5 km SE of Horns, Syria) . 

MATERIAL EXAMINED (2). BM 1965.689 $ (?) Qasr Amra (hamada) ; BM 1965.690 
$ (?) Shishan (hamada) ; both, April-May 1965, S. Bisserot. 



LIZARDS AND SNAKES FROM TRANSJORDAN 237 

PHOLIDOSIS. Scales across middle of body: 60; 62. Longitudinal series of 
ventral plates: 10. Plates in collar: 9. Gular scales in a straight median series: 29; 
28. Femoral pores: 15, 14-15. Lamellae under 4th toe: 20; 24. Upper labials 
anterior to centre of eye: 5, the 5th being the first of two small false supralabials 
below the subocular. 

MEASUREMENTS. SV 46; 49 mm. Tail : 69 (tip missing) ; 81 mm. 
COLORATION. Both pale, the ocelli inconspicuous. 

FIELD NOTES. "The blown sand areas in wadis were the chief habitat of the 
fringe-toed lizards, Acanthodactylus . . . however the lizard (Eremias brevirostris) 
was also seen in this habitat but not as frequently as on the hamada" (S. Bisserot). 

REMARKS. Specimens from eastern and north-eastern Syria, like those from 
Iraq, are so far inseparable from the typical form (Angel, 1936 ; Schmidt, 1939 ; Haas 
and Werner, 1969). Angel's microlepis from western Syria (and a greater altitude) 
had been based on a single specimen, and was not regarded as valid by Haas (1957 : 
73). However, the present two specimens from northern Trans Jordan agree fairly 
well with Angel's description. Furthermore, 13 specimens from central Trans- 
jordan (Amman-Ma'an) in the collection of the Hebrew University (Haas, 1943 : 14) 
show a clear affinity to microlepis, having 46-57 (commonest numbers 53-54) scales 
across the middle of the body, and also relatively small gular scales (21-29, usually 
25-27, in a straight series). Interestingly a specimen from southern Trans Jordan 
(Guweira-Aqaba, HUJ-R 1230) has only 44 scales across the middle of the body (and 
25 gulars). For comparison, Angel's (1936 : 112) E. b. brevirostris from NE Syria 
had 40-52 (commonest numbers 47-49) scales across the middle of the body, and 
20-25, usually 21-23, gular scales in a straight series. Thus E. b. microlepis occupies 
the centre (around Jebel ed Druze) of the western distributional frontier of the species 
(Hoofien, 1957), possibly intergrading with the typical form to the north, east and 
south. 



Eremias guttulata guttulata Lichtenstein 

Lacerta guttulata Lichtenstein, 1823. Verz. Doubl. Mus. Berl., p. 101 (Egypt). 

Eremias guttulata, A. Smith, 1845. 111. Zool.. S. Afr., Kept., PI. 48, fig. 8. 

Eremias guttulata forma typica, Boulenger, 1921. Monograph of the Lacertidae, London, 2, 

p. 258. 
Eremias guttulata guttulata, Wettstein, 1928. Sitzber. Akad. Wiss. Wien (math.-natur.) 137, 

Abt. i, p. 782. 

MATERIAL EXAMINED (i). BM 1965.688 $ 2 miles S of Azraq Druze (basalt 
desert), April-May 1965, S. Bisserot. 

PHOLIDOSIS. Scales across middle of body: 50. Longitudinal series of ventral 
plates : 8. Femoral pores : 13-14. Lamellae under 4th toe : 22-24. Upper labials 
preceding subocular (which enters lip) : 4. 

MEASUREMENTS. SV: 46 mm. Tail: 83 mm (tip regenerated). 



238 Y. L. WERNER 

COLORATION. The dark borders of the dorsal 'ocelli' are black and tend to merge 
with their neighbours laterally, forming incomplete black crossbands, which are 
interrupted by the white centres of the 'ocelli'. 

REMARKS. The snout is very elongated, pointed and flattened. In comparison 
with specimens from Cis Jordan the pileus is smooth and flat, eyes and nostrils being 
little elevated. 

Fourteen other specimens from various localities between Amman and Petra 
(HUJ-R 1237, 1240, 1256-7, 1259-60, 1262-3, 6236-8, 6273-4, 6300) have the 
following ranges of counts: Scales across middle of body, 44-57 (48-52 in 9 speci- 
mens). Longitudinal series of ventral plates, 10. Femoral pores, 10-14 ( I2 ~ I 3 m 
9 specimens). Lamellae under 4th toe, 18-24 ( 22 m 6 specimens). Upper labials 
preceding subocular, 4. The pileus, as in the Basalt Desert specimen, is relatively 
smooth and flat, although in a few specimens the nostrils (and sometimes the eyes 
too) are somewhat elevated. 

The 3 largest specimens measure 50-51 mm (SV). 

In none of these 14 specimens does the dark component of the pattern occupy 
such a large area as in the specimen from the Basalt Desert, nor is this component 
black. In some specimens it is brown, in others, pale to the point of becoming 
indistinct. Some specimens show a tendency for lateral confluence of 'ocelli', but 
the resulting pattern resembles strings of beads rather than crossbands of uniform 
width. The range of coloration known from Cisjordan resembles that shown by 
these 14 specimens. Thus the Basalt Desert specimen is outstanding in its black 
and extensive dark pattern, perhaps as an adaptation to its habitat. 

Ophisops elegans blanfordi Schmidt 
(Text-figs 6, 7) 

Ophisops blanfordi Schmidt, 1939. Zool. Ser. Field Mus. Nat. Hist., 24 (7) : 64-65 (Halfaya, 
20 miles east of Amara, Iraq). 

MATERIAL EXAMINED (2). BM 1963.668 ^ N of Shaubak, 1963, S. Bisserot; BM 
1965.687 $ Ain el Enoquiya (sand and stone wadi), April-May 1965, S. Bisserot. 

PHOLIDOSIS. Scales and plates around middle of body 38; 34. Femoral pores: 
10-11 ; 10-11. Lamellae under 4th toe : 22-24. Upper labials preceding subocular: 
4. Third postsubocular in broad contact with auricular. Postnasal: single. 
Occipital of medium size (somewhat larger than postnasal). 

MEASUREMENTS. SV: 42; 40 mm. Tail of <$: 92 mm; of $ missing. 

COLORATION. Both have the usual Ophisops elegans pattern except that there is 
a distinct dark vertebral line running from the occiput to the pelvic region. 

REMARKS. Both specimens agree with Schmidt's description, except in having a 
slightly higher number of scales around the body (Schmidt's 92 specimens, all from 
the lower Tigris-Euphrates Valley, had 30-36 scales and plates around the middle of 
the body, averaging 33). 



LIZARDS AND SNAKES FROM TRANSJORDAN 239 

The dark vertebral line observed in the two specimens from Transjordan occurs 
only rarely in 0. e. ehrenbergi from Cisjordan (N = 50), and then only on the neck. 
Likewise, Lantz (1930 : 41) says of the pattern of 0. e. elegans "Dessin caracterise 
per la bande occipitale rudimentaire . . . Bande occipitale absente ou reduite a un 
petit trait ou a quelques petites taches noires sur la nuque." It is much commoner, 
and better developed, in 0. e. schlueteri from Cyprus (N = 16). 

Ophisops from Transjordan in the HUJ collection fall into two groups. All 
those from Jerash and its vicinity (N =16: HUJ-R 1190, 1203, 1204/1-2, 1205, 
1206/1-4, 1207/1-3, 1208/1-2, 1209, 1561) have double postnasals. In most of 
them, the vertebral line is either absent, or confined to the occipital region; but in 
3 $3 it extends to the shoulders and in a single male it reaches the midbody although 
it is very faint. These specimens appear to be assignable, like all those from Cis- 
jordan, to 0. e. ehrenbergi. On the other hand, specimens from between Amman 
and Petra are assignable to blanfordi (N =9: HUJ-R 1183, n86, 1218, 1220-22, 
6158-60). Of these, 5 have single postnasals, 3 have double postnasals, and one is 

to 

i/> 

Z 

H- 
I/) 

O n D..O D cm 

oo n v o DO on 



1/2 



01 23456 

VERTEBRAL LINE 

FIG. 6. Ophisops elegans subspp. from. Transjordan. Number of postnasals against extent 
of vertebral line (o, none; 2, only on occiput; 3, reaching shoulder; 4, reaching midbody; 
5, reaching pelvis; 6, reaching tail base. These values are adjusted by + i for unusually 
intense lines, and by -i for particularly faint ones). Open symbols, O.e. ehrenbergi from 
the Jerash area; Solid symbols, O. e. blanfordi (details in text); Squares, males; Circles, 
females; Diamonds, juveniles. 



240 



Y. L. WERNER 



asymmetrical. In most the vertebral line is well developed and this is particularly 
true of those with double postnasals. Thus the two forms are distinguishable by the 
combination of these two characters (Text-fig. 6). Possibly they are also distin- 
guishable by a combination of femoral pore number and vertebral line extent 

(Text-fig. 7). 



a 13 

a. 
to 



to 

UJ 

oe. 

O 

a. 



12 



1 1 



o 10 



D 
D 



D 



a 
oo 



na 



0123456 

VERTEBRAL LINE 

FIG. 7. Ophisops elegans subspp. from Transjordan. Number of femoral pores (repre- 
sented for each specimen as the mean of both femurs) against extent of vertebral line. 
Symbols as in fig. 6. 



The 9 HU J blanfordi specimens have the following scale counts : Scales and plates 
around middle of body, 29-44 (39-41 in 4 specimens) ; Femoral pores, 8-13 ; Lamellae 
under 4th toe, 20-25. Neither these nor the remaining conventional counts differ 
markedly from those found in 0. e. ehrenbergi. 

In conclusion, while the Ophisops of the Jerash district appears to be consub- 
specific with 0. e. ehrenbergi of Cisjordan, the form occurring in the more arid parts 
of Transjordan is Schmidt's 0. blanfordi. This, however, does not seem to merit 
specific rank since there are indications of intergradation with 0. elegans in Trans- 
Jordan (details above) and Iraq (Haas and Werner, 1969), and there is no evidence of 



LIZARDS AND SNAKES FROM TRANSJORDAN 241 

sympatry. Two additional alleged blanfordi characters in fact occur also in (other) 
0. elegans: the small temporal scales, which occur in 0. e. schlueteri (Cyprus) ; and the 
situation of the third post subocular which in 0. e. ehrenbergi (Cisjordan) and 0. e. 
schlueteri sometimes touches and sometimes fails to touch the auricular. 

SCINCIDAE 

Chalcides oceltatus ocellatus Forskal 

Lacerta ocellata Forskal, 1775. Descr. Anim., p. 13 (Egypt). 

Chalcides ocellatus forma typica, Boulenger, 1890. Ann. Mag. Nat. Hist. 5 : 444-445. 
Chalcides ocellatus ocellatus Wettstein, 1928. Sitzber. Akad. Wiss. Wien (math.-natur.), 137, 
Abt. I, p. 784. 

MATERIAL EXAMINED (i). BM 1965.785 Aqaba (on rocks close to beach, Red Sea 
coast), 5 August 1965, D. Western. 

PHOLIDOSIS. Scales around the middle of the body: 30. 
MEASUREMENTS. SV 61 mm; tail: 69 mm. 

COLORATION. Collector's note: "brown with darker brown and white spots". 
The 'ocelli' are numerous and are arranged in transverse series on the neck and tail. 
They are small, each occupying less than a single scale. 

REMARKS. Two out of 3 specimens (the 3rd being damaged) from 65 miles SSE of 
Amman (HUJ-R 1442, 5107, 5108 June-July, 1938 Haas) have 28 scale rows. 
The same applies to HUJ-R 1469 from El-Hamma (NW Transjordan within Israel 
13 March 1945, Haas). The pattern of HUJ-R 5107 is remarkably irregular, many 
of the 'ocelli' having only a black spot on one side of the white centre instead of a 
complete border. In HUJ-R 1442 and 5108 (juveniles) there are only faint indica- 
tions of 'ocelli'. The specimen from El-Hamma has 'normal' 'ocelli', each occupying 
a scale. 

Eumeces schneideri princeps Eichwald 
(PL 5 B) 

Euprepes princeps Eichwald, 1839. Bull. Soc. Imp. Nat. Moscow, 2 : 303-307 ("In ora Caspia 

occidentali, ad montes praesertim Talyschensis") . 
Eumeces princeps, Taylor, 1935. Kansas Univ. Sci. Bull., 23 : 138. 
Eumeces schneideri princeps, Eiselt, 1940. Zool. Anz., 131 : 218. 

MATERIAL EXAMINED (i). BM 1965.695 <$ Ain el Enoquiyya (basalt desert), 
April-May 1965, S. Bisserot. 

PHOLIDOSIS. Scales around middle of body: 27. Dorsal scales from occiput to 
above cloaca: 67. 

MEASUREMENTS. SV: 115 mm (tail regenerated). 

COLORATION. Ground colour of back light brown. No light spots. Along the 



242 Y. L. WERNER 

flank a dark brown band, 2-3 scales broad; its lower border half a scale above the 
light lateral band, its upper border fairly sharp (PI. 5 B). The light lateral band is 
intensely white. (It may have been yellow at the time the animal was killed, six 
months prior to its examination by me.) 

REMARKS. The scale counts of this specimen are characteristic of princeps 
(Eiselt, 1940: Table i), but the colour is unusual, and may represent an adaptation 
to the basalt desert. A specimen from Shaubak (BM 1963.669) here assigned to 
schneideri shows some tendency towards a similar coloration. However, a specimen 
collected 65 miles SSE of Amman (HUJ-R 1389) and clearly assignable to princeps 
(27 scale rows, 69 dorsal scales from occiput to above cloaca) is uniformly coloured 
having neither light spots nor darkened flanks, the light lateral band merging with 
the light belly. 



Eumeces schneideri schneideri Daudin 
(PI. 5 A, C) 

Scincus schneideri, Daudin, 1802. Hist. Nat. Rept., 4 : 291. 

Eumeces schneideri (part*), Taylor, 1935. Kansas Univ. Sci. Bull. 23 : 126. 

Eumeces schneideri schneideri, Eiselt, 1940. Zool. Anz., 131 : 213. 

MATERIAL EXAMINED (2). BM 1963.669 N of Shaubak, 1963, S. Bisserot. BM 
1965.786 Petra (on red sandstone), 1965, D. Western. 

PHOLIDOSIS. Scales around middle of body: 26, 24. Dorsal scales from occiput 
to above cloaca : 65, 66. 

MEASUREMENTS. Largest (BM 1965.786): SV: 114 mm; tail: 209 mm. 

COLORATION. Both specimens have the usual light lateral band passing through 
the ear. BM 1963.669 (PI. 5-C) : Only a few small light (originally yellow-orange ?) 
spots, each covering up to a third of a scale. Flanks mottled dark brown above the 
light band; each dark spot covering the posterior portion of a scale. BM 1965.786 
(PI. 5 A): collector's note: "green with orange spots". The spots each cover up to 
a whole scale, and are irregularly arranged; a tendency to form transverse series is 
particularly evident on the tail. 

REMARKS. These specimens were collected relatively near the area where, in 
Israel, the northwestern pavimentatus, and the southern schneideri intergrade. In 
southern Israel and in Sinai (Schmidt and Marx, 1956 : 28) there occur populations 
in which the pattern is regularly of the schneideri type, but the scale counts of many 
specimens tend towards those characterizing pavimentatus (Eiselt, 1940: Table i). 
This situation is exemplified by BM 1965.786 from Petra, which it seems best to 
assign to schneideri (see also Taylor, 1935 : 130) like the specimens from southern- 

*Taylor includes in schneideri single specimens from "Haiffa" and "Mt. Jerusalem", within the range 
of pavimentatus. These specimens evidently are adult males of pavimentatus which, unlike the females, 
lose the whitish streaks adorning the young. The orange spots however, remain arranged in longi- 
tudinal rows (see his Plate 5). 



LIZARDS AND SNAKES FROM TRANSJORDAN 243 

most Israel. BM 1963.669 from North of Shaubak has 26 scale rows, and its very 
broad dorsal scales exclude it from princeps. 



Mabuya vittata Olivier 

Scincus vittatus Olivier, 1804. Voy. Emp. Ottoman, 3, p. 103, pi. 29, fig. i (sands west of 

Rosetta) . 
Mabuia vittata Boulenger, 1887. Cat. Lizards Brit. Mus., 3, p. 176. 

MATERIAL EXAMINED (i). BM 1963.670 2 km SE Druze village, Azraq, 1963, 
S. Bisserot. 

PHOLIDOSIS. Scales around middle of body: 32. 
MEASUREMENTS. SV: 73 mm; tail: 80 mm. 

COLORATION. Dorsum brown, with three light longitudinal bands. No darker 
spots, except tiny ones on occiput. 

REMARKS. Of 4 specimens from the surroundings of Jerash (HUJ-R 1423, 1424, 
1426, 1547; November 1945, Coll. Haas and Hoofien), 3 have 32 scales around the 
middle of the body, and one has 34. The largest of these measures 95 mm (SV). 
The pattern varies: one specimen resembles BM 1963.670, but another has 5 light 
bands, and two have 4, the median one being obliterated. The other 3 specimens 
have most dorsal scales partly edged in black (or dark brown), particularly towards 
the borders of the light bands. In Cisjordan, too, the pattern and colour of this 
species are highly variable (cf. Peracca, 1894 : 8). 



OPHIDIA 
COLUBRIDAE 

Natrix tessellata tessellata Laurent! 

Coronella tessellata Laurenti, 1768. Synops. Kept.: 87 ("in Japidia, vulgo Cars"). 
Natrix tessellata, Bonaparte, 1834. Iconogr. Faun. Ital., 2, n : plate. 
Natrix tessellata tessellata, (Hecht) 1930. Mitt. zool. Mus. Berlin, 16 : 319. 

MATERIAL EXAMINED (i). BM 1965.696 juv., Shishan (sandy area nr. date palms), 
1965, S. Bisserot. 

PHOLIDOSIS. Scale rows: 19. Ventrals: 165. Subcaudals: 62. 
MEASUREMENTS. SV: 190 mm; tail: 45 mm. 

REMARKS. Three juveniles from Birketen near Jerash (HUJ-R 3024, 3063, 3071 ; 
November 1945, Coll. Haas and Hoofien) have 164-166 ventrals and 65-67 sub- 
caudals. In Cisjordan (N =9), similarly, 160-169 ventrals and 56-66 subcaudals 
have been counted. 



244 Y. L. WERNER 

Coluber rhodorhachis rhodorhachis Jan 

(PI. 6 A, B) 

Zamenis rhodorhachis Jan, 1865. In De Filippi, Viagg. in Persia, p. 356 (Iran; restricted by 

Kramer and Schnurrenberger, 1963, p. 501, to Schiras, Central Persia.) 
Coluber rhodorhachis, Parker, 1931. Ann. Mag. Nat. Hist., (10), 8 : 516. 

Coluber rhodorhachis rhodorhachis, Khalaf, 1959. Reptiles of Iraq with some notes on the 
Amphibians, Baghdad, pp. 75-76. 

MATERIAL EXAMINED (i). BM 1965.805 $ Petra, 1965, D. Western. 
PHOLIDOSIS. Scale rows: 19. Ventrals: 242. Subcaudals: 133. 
MEASUREMENTS. SV: 790 mm; tail: 320 mm. 

COLORATION. Dark crossbands on anterior part of back (65-70) somewhat 
irregular, nearly four times as broad as the light intervening spaces. First dark 
crossband (on the occiput) interrupted mid-dorsally by a faint light vertebral line 
(PI. 6 A). 

REMARKS. This is apparently the first formal record of C. rhodorhachis from 
Transjordan proper. The range extension involved is only minor, as the species is 
well known in the Wadi 'Arava (Haas, 1951). 

Although this specimen was collected at Petra, and its pholidosis agrees with 
material from the adjacent territory of Cisjordan, its coloration deviates markedly 
from that found in these specimens. In Cisjordan the dark crossbands are usually 
distinctly narrower than the intervening light spaces (PI. 6 B), or at the most as 
broad as the latter. The same appears to be true of Egyptian specimens (Anderson, 
1898: pi. 35). On the other hand, I have seen broad and close dark crossbands, like 
those of the Petra specimen, in an example from Iran (MCZ 58872), though in this 
case only the anterior 30-35 crossbands are so broad, the more posterior ones 
gradually become narrower. The similarity of these two specimens does not, 
however, mean very much, as the species exhibits high variability of colour and 
pattern. 

Terent'ev and Chernov (1949 : 242) accept C. r. ladacensis Anderson 1871 
(Boulenger, 1890 : 326) as a distinct form. Mertens (1956 : 95) and Kramer and 
Schnurrenberger (1963 : 501) doubt its validity. In fact, Anderson himself (1895 : 
654, footnote i) says "I am indebted to the Trustees of the Indian Museum for the 
opportunity of re-examining the types of Z. ladacensis. They are unquestionably 
identical with Jan's Z. rhodorhachis. At the time I described the species, Jan's 
work was not in the library of the Indian Museum, Calcutta". Thus, so far all 
Coluber rhodorhachis specimens, apart from the Somalian C. r. subnigra Boettger 
1893 (Parker, 1949 : 30-37), are assignable to the typical form. 

Coluber rogersi Anderson 
(PI. 6 C, D) 

Zamenis rogersi Anderson, 1893. Ann. Mag. Nat. Hist. (6)12 : 439 (Desert to the east of 

Helwan, near Cairo). 
Coluber rogersi, Flower, 1933. Proc. Zool. Soc. London, p. 810-811. 



LIZARDS AND SNAKES FROM TRANSJORDAN 245 

MATERIAL EXAMINED (2). BM 1965.698 $ Five km S of Aseikhim, April-May 
1965, S. Bisserot. BM 1965.806 juv. $ (?) Azraq, 1965, D. Western. 

PHOLIDOSIS. Scale rows: 19; 19. Ventrals: 195; 204. Anals divided. Tails 
incomplete. 

MEASUREMENTS. SV: 565; 220 mm. (Tails incomplete.) 

COLORATION. In both specimens, the anterior three-quarters of body has about 
55 dark, closely set, dorsal blotches (PI. 6 C). Posteriorly the blotches gradually 
become indistinct. The first three blotches are confluent mid-dorsally to form a 
longitudinal streak behind the occiput. Tail uniformly grey. 

REMARKS. The pattern resembles that normally encountered in (southern) 
Cisjordan (PI. 6 D), the blotches being closer to each other than those figured by 
Anderson (1898: pi. 36) for a male from "Beltim" (Nile delta). The scale count 
appears to be higher in Transjordan than in Cisjordan: Two specimens from 65 km 
SSE of Amman have 200 ventrals each (HUJ-R 3183/1 and 2) ; one from N of Zerka 
has 195. Thus while in Cisjordan we find 188-200 ventrals (N = 13), we find 
195-204 (N = 5) in Transjordan. A single specimen from Iraq has 206 (HUJ-R 
3540). 

Malpolon moilensis Reuss 

Coluber moilensis Reuss, 1834. Mus. Senckenb. i, p. 142 (near Moila, on the Red Sea coast of 

Arabia) . 
Malpolon moilensis, Parker, 1931. Ann. Mag. Nat. Hist. (10) 8 : 522. 

MATERIAL EXAMINED (i). BM 1965.697 <$ (?). Three miles N of Druze (Track 
through hamada), April-May 1965, S. Bisserot. 

PHOLIDOSIS. Scale rows: 17. Ventrals: 166. Anal divided. Subcaudals: 50. 
MEASUREMENTS. SV: 443 mm; tail: 95 mm. 

COLORATION (After preservation). Brown with darker brown spots, of varying 
distinctness, which are arranged in eight longitudinal rows. Neighbouring spots 
tend to merge, forming obliquely transverse streaks. A conspicuous dark brown 
blotch is present on each temporal region. 

REMARKS. This appears to be the first record of this snake from Transjordan. 
The species has recently been recorded from southern Cisjordan, where it is rare 
(Barash & Hoofien, 1956). Most, if not all, specimens were actually found in the 
Wadi 'Arava, so that the inclusion of this species in the fauna of Transjordan would 
have been a matter of course. The locality recorded here, however, constitutes a 
significant range extension. 

Malpolon monspessulanus insignitus Geoffroy 

Coluber insignitus Geoflroy in Savignyi, 1827. Descr. Egypte., Hist, nat., i Rept. : 151; pi. 7, 

Fig. 6 (Lower Egypt). 
Malpolon monspessulanus insignitus, Mertens and Miiller, 1928, Abh. senckenberg. naturf. Ges., 

Frankfurt a.M., 41 : 51. 



246 Y. L. WERNER 

MATERIAL EXAMINED (i). BM 1965.807 <$ Azraq, 1965, D. Western. 
PHOLIDOSIS. Scale rows: 19. Ventrals: 173. Anal divided. Subcaudals: 83. 
MEASUREMENTS. SV: 745 mm. Tail: 210 mm. 

COLORATION (After preservation). Nearly uniform dark grey. Belly whitish 
with some light grey mottling. 

DISCUSSION 

The material reported here does not on its own permit an analysis of the herpeto- 
fauna of Trans Jordan in terms of ecology or zoogeography. However, the specimens 
have been assembled on several trips, so that at least the lizard material probably 
reflects to some extent the abundance of the species in certain habitats and localities. 
It thus seems profitable to relate the data available to existing general information 
on the ecological and biogeographical subdivision of Trans Jordan and to the known 
circumstances in Cisjordan. 

General Biogeography of Trans Jordan 

The variegated zoogeography of Transjordan was outlined by Bodenheimer 
(1935 : 24). His basically correct map (1935: Fig. 6) was superseded by the more 
recent work of botanists (Bodenheimer, 1953 : 85). Feinbrun and Zohary prepared 
a phytogeographical map (1955 : 15) dividing Transjordan into the same three 
territories, Mediterranean, Irano-Turanian, and Saharo-Sindian, into which Cis- 
jordan is also divided (Zohary, 1955). This division, based on the mapping of plant 
associations (Feinbrun and Zohary, 1955 : folding map), is related to the distribution 
of soil types and, more closely, to that of rainfall (maps, Feinbrun and Zohary, 1955 : 
9 and 13; Poore and Robertson, 1964 : 12; Bender, 1968 : 10 and 180). The phyto- 
geographical map of Feinbrun and Zohary (1955 : 15) furnished the basis for the 
delimitation of biogeographical territories in Text-fig. 8. (A fourth territory, the 
'Sudanian Penetration Territory', has been proposed for the Lower Jordan Valley 
and the Wadi 'Arava by Gruenberg-Fertig, 1965). The general ranges of the three 
biogeographical territories in southwestern Asia are presented by a map recently 
published elsewhere (Haas and Werner, 1969 : 368). 

Poore and Robertson (1964 : 14-15) similarly classified the Transjordanian range 
types into three basic regions : Mediterranean, Steppe, and Desert of varying type 
(limestone, basalt, sandstone and granite deserts). The salient differences between 
the two maps are few : The Mediterranean region of Poore and Robertson is narrower 
(in W-E direction) than that of Feinbrun and Zohary (shown here in Text-fig. 8) so 
that for instance, Petra, Tafila and Shaubak are excluded from it (the first lying 
just west of, the two last, just east of, Poore and Robertson's Mediterranean region). 
The Steppe region of Poore and Robertson, roughly coinciding in the south with 
Feinbrun and Zohary's Irano-Turanian territory, reaches in the north eastwards to 
only halfway between Amman and Azraq. (Guest (1966: Figs 13, 14) apparently 
includes the area around, and east of, Azraq, in the Irano-Turanian, in agreement 



LIZARDS AND SNAKES FROM TRANSJORDAN 



247 




FIG. 8. The biogeography of Transjordan, based on Feinbrun and Zohary (1955). The 
100 mm mean annual precipitation line is based on Poore and Robertson (1963) ; its inter- 
rupted part (extrapolated alternatives in area lacking data) is based on various sources. 
M, Mediterranean; IT, Irano-Turanian ; SS, Saharo-Sindian. 



248 Y. L. WERNER 

with Feinbrun and Zohary.) At the latitude of the Dead Sea Poore and Robertson's 
Steppe region has a south-eastern extension (as compared to Feinbrun and Zohary's 
Irano-Turanian territory) so as to include El Qatrane. The desert regions of Poore 
and Robertson include areas excluded from Feinbrun and Zohary's Saharo-Sindian 
territory. Firstly, the district around, and east of, Azraq ; secondly, a broader zone 
in the Jordan Valley and especially in its northern part. Thus, Salt (NW of Amman) 
and Petra are well within the Mediterranean territory of Feinbrun and Zohary, but 
on the fringe of the desert according to Poore and Robertson. 

The bioclimates of Transjordan have been defined and mapped by Long (1957; 
also in Poore and Robertson, 1964 : 10, and 13). His map corresponds closely to the 
two biogeographical maps just discussed; concerning the district surrounding, and 
east of, Azraq, it is intermediate between them. The climates of the Mediterranean 
territory are "sub-humid and semi-arid mediterranean bioclimates"; those of the 
Irano-Turanian (~ Steppe) territory are "arid mediterranean bioclimates"; and the 
Saharo-Sindian (~ desert) territory has "saharan mediterranean bioclimates". 
Among the latter, the "cool variety" characterizes the Azraq district. Long's map 
differs from the phytogeographical maps chiefly in that his "semi-arid mediterranean 
bioclimate" (~ Mediterranean territory) reaches southwards only to a point between 
Shaubak and Petra. However, this difference may conceivably result from the 
paucity of meteorological information available to Long (Poore and Robertson, 
1964 : n). 

Thus Transjordan comprises three gross ecological, hence biogeographical terri- 
tories. Relying on the sources cited, these may be briefly characterized as follows : 

The Mediterranean territory includes the hills of Gilead and the western parts of 
Ammon, Moab and N Edom. In the south this narrow territory is confined to the 
higher hills, and interrupted between them. Predominant soils are terra-rossa, 
white Cenomanian soil and grey Senonian soil; along the Jordan Valley, areas of 
Nubian sandstone are included. The bioclimate conforms in the main to the semi- 
arid mediterranean type. Annual rainfall varies from about 700 mm on northern 
mountaintops down to ca. 300, and even below 200, south of Petra. Vegetation is 
characterized by a climax of Mediterranean Maquis and forest types, and by areas 
covered with various shrubs, including many Labiatae. 

The Irano-Turanian territory surrounds the Mediterranean territory except in 
the north (where the latter continues into Syria and Lebanon). Above latitude 
31 30' N the otherwise narrow zone widens eastwards, extending to at least halfway 
between Amman and Azraq, possibly to the frontier. The commonest soils are loess 
and grey calcareous steppe soils (Basalt from Azraq to Burqu). The bioclimate is of 
the arid mediterranean types (of the saharan mediterranean type, cool variety, 
around Azraq) ; annual rainfall is between ca. 350 mm and ca. 100 mm. Vegetation 
is typically a steppe of dwarf -shrubs (Artemisia herba-alba is prominent) or of herbs 
(e.g. Poa sinica and Car ex pachystylis) , and includes remnants of a Pistacia atlantica 
forest. 

The Saharo-Sindian territory occupies the vast southern and eastern desert of 
Transjordan, as well as the lower parts of the Rift Valley. The greatest part, north 
of latitude 29 40' N, is characterized by lime-stone hamadas, the ground generally 



LIZARDS AND SNAKES FROM TRANSJORDAN 249 

being covered by a layer of flints. Further south, and in the Rift Valley, Nubian 
sandstone and sand predominate, with an area of granite (rock and sand) in the Wadi 
'Arava. A vast basalt desert, from Azraq to Burqu, is largely covered by basalt 
boulders of varying size. The bioclimates are of the Saharan Mediterranean types. 
Rainfall is normally below 150 mm, in the extreme southeast below 50 mm. Vegeta- 
tion is scanty except in the wadis. Associations of Anabasis articulata are promi- 
nent, and on granite sands Haloxylon spp. 

Distribution of Reptiles 

This discussion is limited to the 23 species and subspecies reported here. Among 
these, 16 occur also in Cisjordan. Two ubiquitous species were collected in all three 
regions of Transjordan (and similarly occur in all parts of Cisjordan) : Hemidactylus 
turcicus and Chalcides o. ocellatus. These have an essentially circum-mediterranean 
distribution, which is particularly broad to the south and east. Agama stellio is 
similarly circum-eastern-mediterranean, but unfortunately its infraspecific taxonomy 
is not clear. 

Four forms occur in Cisjordan as Mediterranean elements: Chamaeleo chamaeleon 
recticrista, Mabuya vittata, Natrix t. tessellata and Malpolon m. insignitus. The first 
three are here reported from the Mediterranean territory of Transjordan; Mabuya 
vittata and Natrix t. tessellata were also found at Azraq or at the neighbouring 
Shishan (within the disputed area belonging to either the Irano-Turanian or Saharo- 
Sindian territory). Malpolon m. insignitus is reported from Azraq. This distribu- 
tion is probably due to the local conditions prevailing at these places. N. t. tessellata 
is semiaquatic; M. vittata is facultatively hydrophilic, and lives, e.g., also among 
reeds surrounding the salt marshes south of the Dead Sea (pers. obs.). 

Seven reptiles typical of Cisjordanian desert habitats were collected in the Irano- 
Turanian and Saharo-Sindian areas of Transjordan: Agama sinaita, Acanthodactylus 
b. asper, Eremias g. gutulata, Eumeces s. schneideri, Coluber rhodorhachis, C. rogersi, 
and Malpolon moilensis. We shall consider first the two Coluber species. All of 4 
localities for Coluber rogersi fall within the Irano-Turanian (according to Feinbrun 
and Zohary; but 2 within the desert according to Poore and Robertson.). In Cis- 
jordan this snake occupies both the Irano-Turanian and Saharo-Sindian territories, 
with the exception of the (hot) Wadi 'Arava. Its world distribution is Saharo- 
Sindian and to some extent, Irano-Turanian. C. rhodorhachis occurs in Cisjordan 
mainly in the Saharo-Sindian territory (Wadi 'Arava and southern Negev), and the 
first and only record for Transjordan is from Petra, on the fringe of the Wadi 'Arava. 
Its world distribution is Irano-Turanian and Saharo-Sindian. The two lacertids, 
A. b. asper and E. g. guttulata, are reported from both Irano-Turanian and Saharo- 
Sindian localities in Transjordan. Similarly, they are known in Cisjordan from 
suitable habitats throughout the Negev. Their world distribution is Saharo- 
Sindian. The remaining forms, Agama sinaita, Eumeces s. schneideri and Malpolon 
moilensis, were taken in Transjordan at more or less desertic localities (at least 
according to Poore and Robertson). In Cisjordan these are restricted to the 
Saharo-Sindian part of the Negev, and their world distribution is Saharo-Sindian. 



250 Y. L. WERNER 

The ranges of the two forms of Ptyodactylus apparently fail to coincide with any of 
the ecological territories described above. P. h. puiseuxi, a morphologically well 
denned form, is common in northernmost Cis Jordan and in Trans Jordan, on the hills 
bordering Lake Tiberias on the east, and around Jerash all in the mediterranean 
territory but is here reported also from the fringe of the basalt desert ca. lokm E 
of Azraq (Wadi Aseikhim). It may eventually turn out to be basically a form 
inhabiting basalt rocks, which has spread to adjacent habitats. P. h. guttatus is less 
well defined and in particular not clearly distinguishable from P. h. hasselquistii. 
Geckos currently assigned to guttatus are common in the Mediterranean, Irano- 
Turanian and Saharo-Sindian territories of Cisjordan in effect throughout the 
country excepting its northern and southern extremes. So far, our records for 
Transjordan are all within the (southwestern) Saharo-Sindian (Rum; S of Guweira) 
or on its rim (Petra; Wadi Musa). Evidently the ranges of these two geckos are 
influenced to a great extent by factors other than the climate, perhaps because they 
can modify their exposure to it by varying the relative duration of nocturnal and 
diurnal activity. 

Of the Transjordanian reptiles reported here, seven forms do not occur in Cisjordan. 
Of these, six are apparently Irano-Turanian elements, as far as their general ranges 
are concerned: Agama p. haasi, Acanthodactylus grandis, A. t. tristrami, Eremias b. 
microlepis, Ophisops e. blanfordi, and Eumeces s. princeps. In Transjordan, A. p. 
haasi has been collected mostly within the Irano-Turanian, but also within the 
Saharo-Sindian. In Cisjordan, A. p. pallida similarly occurs in both territories. 
A. grandis has been collected in the Irano-Turanian and Saharo-Sindian, and A. t. 
tristrami in the Irano-Turanian and Mediterranean. Some of the localities for E. b. 
microlepis are within the Irano-Turanian, the rest within the Saharo-Sindian. 
These three lacertids have no conspecific relations in Cisjordan. 0. e. blanfordi has 
been collected within the Irano-Turanian and on both its mesic and desertic borders. 
The related 0. e. ehrenbergi occurs in the Mediterranean and Irano-Turanian of 
Cisjordan, and also in the Mediterranean of Transjordan (around Jerash). 

The case of ". s. princeps is of particular interest since three distinct, apparently 
allopatric, races of E. schneideri occur in Transjordan and adjacent areas (Mertens, 
1920, 1924, 1946; Taylor, 1935 ; Eiselt, 1940). Both localities reported here for E. s. 
princeps are within the Irano-Turanian, and this agrees with this race's general 
distribution. In the Saharo-Sindian of both Cis- and Transjordan E. s. schneideri 
occurs, while E. s. pavimentatus lives in the Mediterranean of Cisjordan. Its 
occurrence in Transjordan, which is probable, remains to be shown. 

The last of the Transjordanian reptiles not occurring in Cisjordan is Agama 
blanfordi fieldi (A. persica fieldi Haas and Werner, 1969). This obviously is a 
Saharo-Sindian, Arabian, form. Its taxonomic relationship with the superficially 
similar psammophile, A. savignii of eastern Egypt, Sinai and southern Cisjordan, 
has not been studied but the two probably occupy comparable ecological niches. 

In conclusion, the locality data presented here for Transjordanian lizards and 
snakes are in good agreement with a generalized subdivision of Transjordan into 
three major ecological-biogeographical territories, based on both the maps of Fein- 
brun and Zohary (1955 : 15) and Poore and Robertson (1964 : 14-15). The species 



LIZARDS AND SNAKES FROM TRANSJORDAN 251 

of the Mediterranean territory exhibit no marked morphological deviations from 
their conspecific counterparts in the Mediterranean of Cisjordan. On the other hand, 
in several species of desert reptiles (Saharo-Sindian or Saharo-Sindian and Irano- 
Turanian), the Transjordan population appears to differ from the Cisjordanian one. 
Furthermore, one Saharo-Sindian species, Agama Uanfordi, does not occur west of 
the Rift Valley. Conspicuous is the occurrence in Transjordan of six Irano- 
Turanian species, of which three do not occur in Cisjordan, while the remaining three 
are represented there by other (well defined) subspecies (see also Haas, 1952). The 
reciprocal phenomenon also exists as not all Cisjordanian reptiles occur in Trans- 
Jordan. Thus among the Saharan psammophile reptiles of southern Cisjordan, five 
occur only west of the Rift Valley. Another species, Sphenops sepsoides, penetrates 
into the Rift Valley (Werner, 1968), and only one other, Acanthodactylus scutellatus 
scutellatus, is represented east of the Rift Valley by another subspecies, A. s. hardyi 
(northwestern Saudi Arabia Haas, 1957; Iraq Haas and Werner, 1969). It is 
tempting to assume that the Wadi 'Arava, together with the very steep mountain 
slopes bordering it on the east, constitutes a barrier to the distribution of reptiles, 
though more direct evidence on this effect would be desirable. 



ACKNOWLEDGEMENTS 

For the privilege of examining this material I am grateful to A. G. C. Grandison, 
British Museum (Nat. Hist.). I am much obliged to S. Bisserot who kindly presented 
his field notes for inclusion in this report; to D. Western for his cooperation and 
stimulating correspondence; and to K. Klemmer, Senckenberg Museum, who most 
helpfully supplied information on the type of Agama pallida Reuss, photographs of 
it, and a copy of its description. 

I wish to thank S. Furman and N. Shulman for assistance in examining specimens; 
E. Alcalay and T. Sheffer for help in preparing the diagrams and maps; E. Haupt 
for the photographs of Agama pallida Reuss (type) ; E. Ben-Hur and A. B. Niv for 
the other photographs; and my wife, Nurit, for both advising me on statistics and 
performing the calculations. 

I am indebted to E. N. Arnold, British Museum (Nat. Hist.), G. Haas and, 
particularly, to J. H. Hoofien, for their painstaking and instructive comments on 
the text and illustrations, most of which I heeded. 



APPENDIX 

LIST OF LIZARDS AND SNAKES SO FAR RECORDED FROM 
TRANSJORDAN (INCL. WADI 'ARAVA) 

The documentation cited in parentheses, for species not represented in this report, 
is not necessarily the earliest one available. The names listed are not necessarily 
those employed for the same taxa by the authors cited. 



252 Y. L. WERNER 

LACERTILIA 
GEKKONIDAE 

1. Bunopus blanfordii Strauch (Barash and Hoofien, 1956) 

2. Ceramodactylus doriae Blanford (Haas, 1956) 

3. Hemidactylus turcicus turcicus L. 

4. Pristurus flavipunctatus guweirensis Haas (Haas, 1951) 

5. Ptyodactylus hasselquistii guttatus von Heyden 

6. Ptyodactylus hasselquistii puiseuxi Boutan 

7. Stenodactylus grandiceps Haas (Haas, 1951) 

8. Stenodactylus sthenodactylus sthenodactylus Lichtenstein (Haas, 1951) 

9. Tropiocolotes steudneri Peters (Haas, 1951) 

AGAMIDAE 

10. Agama blanfordi fieldi Haas and Werner 

11. Agama pallida haasi Werner 

12. Agama sinaita von Heyden 

13. Agama stellio brachydactyla Haas 

14. Agama stellio picea Parker (Haas, 1951) 

15. Agama stellio stellio L. (Daan, 1967) 

CHAMAELEONIDAE 

16. Chamaeleo chamaeleon recticrista Boettger 

LACERTIDAE 

17. Acanthodactylus boskianus asper Audouin 

18. Acanthodactylus cantoris schmidti Haas (Hoofien, 1965) 

19. Acanthodactylus grandis Boulenger 

20. Acanthodactylus robustus Werner (Haas, 1951) 

21. Acanthodactylus tristrami tristrami Giinther 

22. Eremias brevirostris microlepis Angel 

23. Eremias guttulata guttulata Lichtenstein 

24. Eremias guttulata watsonana Stoliczka? (Wettstein, 1951) 

25. Eremias olivieri schmidti Haas, (Haas 1951) 

26. Lacerta danfordi danfordi Giinther (Hoofien, 1969) 

27. Ophisops elegans blanfordi Schmidt 

28. Ophisops elegans ehrenbergi Wiegmann (Haas, 1951) 

29. Ophisops elegans elegans Menetrie's (Schmidt, 1939) 

SCINCIDAE 

30. Ablepharus kitaibelii Bibron et Bory (Haas, 1951) 

31. Chalcides ocellatus ocellatus Forskal 

32. Eumeces schneideri princeps Eichwald 



LIZARDS AND SNAKES FROM TRANSJORDAN 253 

33. Eumeces schneideri schneideri Daudin 

34. Mabuya vittata Olivier 

35. Ophiomorus latastii Boulenger (Anderson & Leviton, 1966) 

36. Scincus sp. (Haas, 1951) 

37. Sphenops sepsoides Audouin (Werner, 1968) 



ANGUIDAE 

38. Ophisaurus apodus Pallas (Haas, 1951) 

VARANIDAE 

39. Varanus griseus griseus Daudin (Haas, 1951) 



OPHIDIA 

LEPTOTYPHLOPIDAE 

40. Leptotyphlops phillipsi Barbour (Haas, 1951) 

COLUBRIDAE 

41. Coluber jugularis jugularis L. (Haas, 1951) 

42. Coluber najadum Eichwald (Haas, 1951) 

43. Coluber ravergieri nummifer Reuss (Flower, 1933) 

44. Coluber rhodorhachis Jan 

45. Coluber rogersi Anderson 

46. Eirenis collaris Me'ne'trie's (Haas, 1951) 

47. Eirenis coronella coronella Schlegel (Schmidt, 1939) 

48. Eirenis coronella fraseri Schmidt (Haas, 1951) 

49. Eirenis decemlineata Dumeril & Bibron (Haas, 1951) 

50. Eirenis rothi Jan (Haas, 1951) 

51. Malpolon moilensis Reuss 

52. Malpolon monspessulanus insignitus Geoffroy 

53. Natrix tessellata tessellata Laurenti 

54. Psammophis schokari Forskal (Haas, 1951) 

55. Rhynchocalamus melanocephalus Jan (Hart, 1891) 

56. Spalerosophis diadema ssp. (Haas, 1951) 

57. Tarbophis nigriceps Ahl (Haas, 1951) 

VIPERIDAE 

58. Cerastes cerastes L. (Haas, 1951) 

59. Echis colorata Gunther (Haas, 1951) 

60. Pseudocerastes fieldi Schmidt (Haas, 1951) 



254 Y. L. WERNER 

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Dr. Y. L. WERNER 

Department of Zoology 

HEBREW UNIVERSITY OF JERUSALEM 

JERUSALEM, ISRAEL 




PLATE i 

Agama pallida subspp. : 

(A-D) A. p. haasi subsp. nov. : (A) Holotype^, BM 1965.800, dorsal view; (B) Same, left ear; 
(C) Juvenile, BM 1965.797, right ear; (D) Paratype $, HUJ-R 1884. 

(E-F) A. p. pallida Reuss : (E) From southern Cisjordan (Wadi Ajram) <$, HUJ-R 1623; 
(F) From eastern Egypt (Kassassin) $, HUJ-R 1126. 

Scale, cm and mm (D-F at same magnification). 



Bull. Br. Mus. nat. Hist. (Zool.) 21,6 



PLATE i 



0123 



CH & 8 L 





PLATE 2 

Agama pallida pallida Reuss holotype $, SMF 10007: 
(A) Dorsal view; (B) Head. 



Bull. Br. Mus. not. Hist. (Zool.) 21, 6 



PLATE 2 





PLATE 3 

(A) Agama blanfordi fieldi $ , BM 1965.686. 

(B-E) Agama sinaita: (B) Dorsal view of left pes, and (C) of base of tail, of ^ from northern 
Transjordan (Wadi Ratam), BM 1965.685; (D) Dorsal view of left pes of $ from southern 
Cisjordan, HUJ-R 1919; (E) Dorsal view of tail base of <$ from southern Cisjordan, HUJ-R 1794. 
Scale, cm and mm (B-E at same magnification). 



Bull. Br. Mus. nat. Hist. (Zool.) 21,6 



PLATE 3 







PLATE 



(A-B) Acanthodactylus grandis : (A) Adult $ from Tell el Mukheizin, BM 1965.694; (B) 
Young $ from Ain el Enoquiyya, BM 1965.692. 

(C-D) Acanthodactylus tristrami tristrami : (C) $ from the Jordanian-Syrian border, BM 
1962.352; (D) 3 1 from 15 Km S of Amman, HUJ-R 1333. 

Scale, cm and mm (all at same magnification). 



Bull. Br. Mus. nat. Hist. (Zool.) 21,6 



PLATE 4 




PLATE 5 

Eumeces schneideri subspp. : 

(A) E. s. schneideri from Petra, BM 1965.786; (B) E. s. princeps from Ain el Enoquiyya, 
BM 1965.695; (C) E. s. schneideri from N Shaubak, BM 1963.669. 

Scale, cm and mm. 



Bull. Br. Mus. nat. Hist. (Zool.) 21, 6 



PLATE 5 




PLATE 6 

(A-B) Coluber rhodorhachis rhodorhachis: (A) <j> from Petra, BM 1965.805; (B) <$ from southern 
Cisjordan, HUJ-R 3211. 

(C-D) Coluber rogersi : (C) $ from 5km S of Aseikhim, BM 1965.698; (D) 9 from southern 
Cisjordan, HUJ-R 8020. 

Scale, cm and mm. 



Bull. Br. Mus. nat. Hist. (Zool.) 21,6 



PLATE 6 




:j I i II j II i 1 1 1 1 1 1 1 i 1 1 1 
OARISTO1 2 



'& 



A LIST OF SUPPLEMENTS 
TO THE ZOOLOGICAL SERIES 

OF THE BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 



1. KAY, E. ALISON. Marine Molluscs in the Cuming Collection British Museum 
(Natural History) described by William Harper Pease. Pp. 96; 14 Plates. 
1965. (Out of Print.) 3.75. 

2. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and 
Valenciennes. Pp. 180; n Plates, 15 Text-figures. 1967. 4. 

3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure of Mineralogy 
at the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125; 29 Plates, 
77 Text-figures. 1969. 4.50. 

4. HAYNES, J. R. Cardigan Bay recent Foraminifera (Cruises of the R.V. Antur) 
1962-1964. (In press.) 



Printed in England by Staples Printers Limited at their Kettering, Northants, establishment 






i \ CONCHOECIA FROM THE 
NORTH ATLANTIC 

THE 'PROCERA' GROUP 



M. V. ANGEL 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 21 No. 7 

LONDON: 1971 



CONCHOECIA FROM THE NORTH ATLANTIC 
THE 'PROCERA' GROUP 




BY 



MARTIN VIVIAN ANGEL V|J 

/NU 

National Institute of Oceanography 



Pp. 257-283 ; 14 Text-figures 



THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 21 No. 7 

LONDON: 1971 



THE BULLETIN OF THE BRITISH MUSEUM 

(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 21, No. 7 of the Zoological 
series. The abbreviated titles of periodicals cited follow 
those of the World List of Scientific Periodicals. 



World List abbreviation 
Bull. Br. Mus. nat. Hist. (Zool.). 



Trustees of the British Museum (Natural History), 1971 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 3 December, 1971 Price 1.05 



CONCHOECIA FROM THE NORTH ATLANTIC 
THE 'PROCERA' GROUP 

By MARTIN VIVIAN ANGEL 

INTRODUCTION 

MULLER (1894) decided that Conchoecia variabilis Miiller 1890 was a confusion of 
two species, C. oblonga Claus 1890 and a new species C. procera. Claus (1894) 
also confused these two species. Later Muller (i9o6a) further described C, procera, 
and gave its size range as 1-05-1-35 mm for females and 0-85-1-20 mm for males. 
The geographical range was described as being between 3iN and 35 S in all oceans 
(Muller I9o6a, b, 1908, 1912). It has since been further reported in the North- 
eastern Atlantic (Fowler 1909, Granata & Caporiacco 1949, Angel 1968, iQ6ga., b), 
the Sargasso Sea (Deevey 1968), the Benguela Current (lies 1953), western Mediter- 
ranean (Leveau 1965), the Adriatic (Schweiger 1912, Mure" 1955, 1961), the western 
Indian Ocean (Leveau 1967, 1968), the eastern Indian Ocean (McKenzie in press). 

In 1968 R.R.S. 'Discovery' took a series of hauls in the region of nN 20W, 
about 300 miles south of the Cape Verde Islands, to study vertical distribution 
patterns. It was soon apparent that adult specimens attributable to C. procera 
separated into three distinct size groupings, which showed differential depth distri- 
butions; Deevey (1968) also noted two size groupings. The middle sized group, the 
most abundant, is considered here to be synonomous with C. procera sensus strictu. 
The morphological differences between this group and the smaller and larger groups 
are sufficiently great to attribute them with specific status. They are, therefore, 
named Conchoecia microprocera sp. nov. and Conchoecia macroprocera sp. nov. 
respectively. 

A single adult female taken in a deep vertical haul at 'Discovery' station 4768 
in 1961 near 40 N 2OW, is also described and is ascribed to C. vitjazi Rudjakov 
1962, another species which possibly belongs to the 'procera' group. 

Conchoecia procera Muller 1894 

MALE. Carapace. The lengths of 234 specimens from nN 20W ranged from 
0-98-1-04 mm with a mean of 1-020 + 0-019 mm, and of 70 specimens from off the 
Moroccan coast 34N 8W (Angel 1968) ranged from 0-96-1-04 mm with a mean of 
1-005 0-009 mm - The outline of the carapace (fig. lA, B) with the smooth curve 
of the posterior edge into the ventral edge broken only by the step at the opening 
of the right asymmetrical gland is similar to Miiller's figure (1894 T. XIII fig. 39). 
Similarly the shape of the posterior dorsal spine (fig. 3F) is identical to Miiller's 
figure (1894 T. XIII fig. 41) and is the character by which he separated C. procera 
from C. decipiens Muller in his key (Muller 1912). Edge glands are numerous down 
the posterior carapace margin, and are present but less abundantly along the 



260 



M. V. ANGEL 




K 



FIG. i. Conchoecia procera Muller, male. A, Lateral view of carapace; B, Ventral view 
of carapace; C, Frontal organ and first antenna; D, Armature of the b and e setae of the 
first antenna; E, sixth limb; F, second antenna; G, endopodite of the right second 
antenna; H, left hook appendage; I, right hook appendage; J, penis; K, detail of tip 
of penis. 



CONCHOECIA FROM THE NORTH ATLANTIC 261 

ventral margin. Live animals have not been observed to bioluminesce. 

Frontal organ (fig. iC). The shaft reaches level with the end of the limb of the 
first antenna. The capitulum is downturned and slightly curved. There is a small 
ventral swelling about half way along its length. 

First antenna (fig. iC). The first segment is shorter than the second. None 
of the segments carry any additional armature. The a seta is almost as long as the 
limb, reaching well beyond the joint between the first and second segments. The 
c seta is very short. The remaining three setae are almost equal with the e seta 
just the longest. The d seta is bare but the b seta has three small spines opposite 
the distal end of the e seta armature. The e seta has about seventeen pairs of 
long slender spines (fig. iD) with two pairs of distally pointing spinules just distal 
of the main armature. The main spines decrease in length slightly towards the base 
of the seta. 

Second antenna (fig. iF). The exopodite segments carry no unusual armature. 
The protopodite is a little less than half the carapace length and the first exopodite 
segment is about 3/8's its length. On the endopodite (fig. iG) the processus mamil- 
laris is blunt with a small rounded tubercle slightly offset to one side. The a seta 
which carries fine spinules, is half the length of the b seta. The b seta is armed 
with much longer and stronger spines. The c and d setae are almost as long as the 
second segment, while the e seta is a minute spine. The f seta is only slightly 
shorter than the g seta, and both setae are thin walled terminally and unarmed. 
The h, i and j setae are short with barely developed shafts, but the h seta does have 
a small basal swelling. The left hook appendage (fig. iH) curves through about 
120, terminally tapering into a smooth point. The right hook appendage (fig. iG) 
is larger and slightly more curved. It is slightly swollen and ridged subterminally 
and ends in a point. 

Mandible (fig. 2C). There is only a single long seta on the inner surface of the 
first endopodite segment, and there is no additional armature on any of the other 
endopodite segments. The toothed edge of the basale (fig. 2B) is typical for the 
genus with two tubiform teeth, six serrate teeth and an outer tooth which is unusually 
broad in this species. One of the outer setae is long, but the other only reaches 
level with the teeth. There are long hairs on the outer surface arranged in four rows, 
two leading up to the bases of the spine teeth. The outer teeth have very clear 
secondary tooth rows. The toothed edge of the coxale has ten teeth (fig. 2 A). 
The distal tooth list consists of two large teeth, neither of which is serrate, and ten 
to twelve small teeth, the outermost of which is broad. The proximal list consists 
of 13-15 fairly regular teeth which decrease in size along the list. 

Maxilla (fig. 2D). There is a basal seta. The first endopodite segment has six 
anterior, one lateral and three posterior setae. There is no distal armature on the 
first segment. 

Labrum (fig. 2E). The hyaline membrane has a smooth rather shallow notch. 

Caudal furca (fig. 2G). There is no unpaired seta dorsal to the eight pairs of hook 
spines. There is a covering of fine hairs between the caudal lamellae. 

Penis (figs ij, K). The intromittent organ has five oblique muscles. The end 
of the organ is rounded in outline. 



2&2 



M. V. ANGEL 




FIG. 2. Conchoecia procera Miiller. A, mandible tooth list and toothed edge of the coxale ; 
B, mandible toothed edge of the basale; C, mandible endopodite; D, maxilla endopodite; 
E, labrum; F, fifth limb; G, caudal furca. 



CONCHOECIA FROM THE NORTH ATLANTIC 



263 




B 




FIG. 3. Conchoecia procera Miiller, female. A, lateral view of carapace; B, ventral view 
of carapace; C, frontal organ and first antenna; D, second antenna; E, second antenna 
endopodite; F, male detail of the posterior dorsal corner of the carapace. 



FEMALE. Carapace (figs 3A, B). The lengths of 282 specimens from nN 
ranged from 1-12-1-24 mm with a mean of 1-181 + 0-021 mm. 122 specimens from 
off the Moroccan coast ranged from 1-12-1-24 mm with a mean of 1-166 + 0-022 mm. 
The change in the mean size with latitude is insignificant. 

Frontal organ (fig. 3C). The capitulum is not differentiated from the stalk. 
The total length is about three times the length of the limb of the first antenna It 
has a slight terminal swelling with a rounded knob. There are spines on its under- 
side. 



264 M. V. ANGEL 

First antenna (fig. 36) . There is no additional armature. The a-d setae are 
thin walled and a little longer than the length of the limb. The e seta is nearly 
twice as long as the other setae and distal of their ends it carries long fine spinules 
on the anterior edge and shorter spinules on its trailing edge. 

Second antenna (fig. 3D). The protopodite is 2/5's the length of the carapace 
and nearly three times the length of the first exopodite segment. The longest 
swimming seta is 3/4 the length of the protopodite. On the endopodite (fig. 3E) 
the a seta is bare and less than half the length of the b seta. The b seta carries 
8-9 strong spinules. On the segment near the bases of the a and b setae are about 
eight small spines. The second segment carries many long fine hairs (c.f. Miiller 
1894 T.6 fig. 61). The c, d and e setae are all absent. All the main setae are 
thin walled. The g seta is distinctly broader but only slightly longer than the other 
four subequal setae. 

Synonomy. There is no clear type locality for this species. Miiller (1890) 
described C. variabilis which he later (1894) decided was a confusion between C. 
oblonga Claus and C. procera, from the Pacific and also from the Gulf of Naples. 
Claus's (1894) material described as C. oblonga Claus included C. procera Miiller 
(Miiller I9o6a) came from the eastern Mediterranean. It is not until Miiller's 
(1894) original description of C. procera that any certainty can be attached to the 
identifications. However, there are no localities given for any of the material. 
From Leveau's (1965) rather sparse data and some specimens from the North 
Adriatic sent to me by Professor J. Stirn it has been possible to confirm the presence 
of C. procera sensu strictu in the Mediterranean. Miiller's (igoGa) material included 
at least one if not both of the new species described below, judging from his data 
on size ranges and the armature of the male antennular e seta. Fowler (1909) 
appeared only to have caught C. procera sensu strictu from the Bay of Biscay. 
Deevey (1968) reported two forms of C. procera, the larger of which conformed with 
the description above of C. procera sensu strictu. A re-examination of Angel's 
material from the Moroccan coast (1968) and from near Fuerteventura in the 
Canary Islands (i969a) showed that they all beonged to C. procera sensu strictu, 
with the single exception of the one adult female caught above the thermocline 
at 'Discovery' station 6183 haul 2. This specimen corresponded to the new species 
described below C. microprocera. 

Conchoecia microprocera sp. nov. 

MALE. The holotype mounted on slides in Euparal and stained with lignin 
pink has been deposited in the British Museum (Natural History), No. 1971.2.1.1. 
Locality 'Discovery' station 6665 haul 4, io32.7'N, i9574'W. Depth 400- 
295 m. Time 1559-1731 hrs. Date 22 February 1968. Net Modified Indian 
Ocean Standard Net (Nii3) fitted with a catch dividing bucket (Foxton 1963, 1969). 

Carapace (figs lA, B). The range in length of 204 specimens was 0-82-0-92 mm 
with a mean of 0-863 + 0-013 mm The height and breadth of the carapace were 
both approximately 2/5's the length. In C. procera sensu strictu the carapace 
height was always measurably larger than its breadth (Table i). There was a small 



CONCHOECIA FROM THE NORTH ATLANTIC 



265 




FIG. 4. Conchoecia microprocera sp. nov., male. A, lateral view of carapace; B, ventral 
view of carapace; C, frontal organ and first antenna; D, first antenna armature on e seta; 
E, sixth limb; F, second antenna; G, endopodite of the right second antenna; H, left 
hook appendage; I, penis. 



266 M. V. ANGEL 

spine on the right valve at the posterior dorsal corner (fig. 6F). The asymmetrical 
glands and the edge glands are very similar in their positions to C. procera sensu 
strictu. 

Frontal organ (fig. 46). The stalk projects just beyond the length of the first 
antenna. The capitulum is long and slender and curves slightly downwards. 
Terminally it is rounded. There is a patch of fine long hairs in the centre of the 
ventral surface. 

First antenna (fig. 46). There is no additional armature. The first and second 
segments are subequal, and there are large pigment corpuscles in the region of their 
articulation. The a seta lies back parallel with the limb and reaches almost to the 
base. It is relatively longer than in the other two species (Table i). The c seta 
is extremely short. The b and d setae are subequal, only slightly shorter than the 
e seta. Both carry very fine spinules on a level with the distal end of the e seta 
armature. The e seta armature (fig. 4D) consists of 11-13 pairs of spines which 
increase very slightly in length towards the base of the seta. Beyond these main 
spines are two pairs of distally pointing spinules. Deevey (1968) described some of 
her specimens of C. procera as having this type of armature. 

Second antenna (fig. 4?). The protopodite is more than half the length of the 
carapace, and this distinguishes this species from the others (Table i). The first 
exopodite segment is a third the length of the protopodite and twice the lengths 
of the remaining exopodite segments. The longest swimming seta is about 4/5*5 
the length of the protopodite. There is no additional armature on the exopodite 
segments. On the endopodite (fig. 40) the processus mamillaris has a very bluntly 
pointed tip. The a seta is bare and half the length of the b seta. The b seta carries 
spinules for most of its length. The c and d setae are almost as long as the second 
segment. The e seta is a minute spine. The g and f setae are bare, terminally 
thin walled, blunt and slightly flattened. The h, i and j setae are short with very 
poorly developed shafts. The' left hook appendage (fig. 46) is right angled at its 
base, and then distally straight. It broadens near its end and then tapers to end 
in a smooth curved point. The right hook appendage (fig. 4H) is longer beyond the 
basal angle, and curves to terminate in a curved point with marked subterminal 
ridging. 

Mandible (fig. 56). The toothed edge of the basale is the usual structure for the 
genus (fig. 56). The outer two setae are very short not reaching level with the 
teeth. There are fine hairs on the outer surface, with two rows running up to the 
bases of the tubiform teeth. There are no hairs on the basale between the toothed 
edge and the articulation with the first endopodite segment. On the first endo- 
podite segment there are two setae on the inner edge; one very long, the other 
minute. The terminal claw seta is particularly long. 

The toothed edge of the coxale (fig. 5A) has ten teeth. The distal tooth list con- 
sists of two large teeth, neither serrate, followed by 13-14 small regularly sized 
teeth. The proximal list consists of 15 irregular teeth. 

Maxilla (fig. 5D). There is a basal seta. There are three posterior setae and a 
lateral seta on the first segment, but there are only five anterior setae compared 



CONCHOECIA FROM THE NORTH ATLANTIC 



267 




FIG. 5. Conchoecia microprocera sp. nov. A, mandible tooth lists and toothed edge of 
the coxale; B, mandible toothed list of the basale; C, mandible endopodite; D, maxilla 
endopodite; E, labrum; F, fifth limb; G, caudal furca. 



268 M. V. ANGEL 

with six in the other species of the 'procera' group. There is a row of fine spinules 
on the distal edge of the first endopodite segment. 

Labrum. There is a shallow smooth notch in the hyaline membrane. 

Penis (fig. 4!). There are four oblique muscles. The terminal incurved end plate 
has two small spines. 

Caudal furca (fig. 5G). There is no unpaired seta dorsal of the hook spines. 
There are fine hairs between the two caudal lamellae which are most clearly seen 
between the bases of the first and second hook spines. 

FEMALE. The paratype specimen mounted on slides in Euparal and stained 
with lignin pink has been deposited in the British Museum (Natural History) No. 
1971.2.1.2 The collection data is as for the male. 

Carapace (figs 6A, B). The range in length of 260 specimens was 0-92-1-06 mm 
with a mean of 0-999 0*023 mm - The general appearance was more hyaline 
than for the other closely related species. It is slimmer in appearance, but the 
tendency for the animals to splay open on preservation probably contributed to the 
insignificance in the difference between its relative breadth and those of the other 
species. The openings of the various glands were similar to those of the other 
species. The spine on the right valve at the posterior dorsal corner is well de- 
veloped. 

Frontal organ (fig. 6C). The frontal organ shows some evidence of differentiation 
into stalk and capitulum. The capitulum is a little broader and is swollen towards 
its tip finally ending in a down curved point. 

First antenna (fig. 6C). There is no dorsal seta or other additional armature. 
The a-d setae are only half the length of the e seta. Distal of their ends the e seta 
carries short spinules on its posterior edge and sparse long hairs on its anterior 
edge. 

Second antenna (fig. 6D) . The protopodite is much longer relative to the carapace 
than in the other species (Table i). There is no unusual armature on the exopodite. 
On the endopodite (fig. 6E) the processus mamillaris is rounded. The a seta is 
half the length of the b seta and carries very fine spinules. The b seta has four 
to six stouter spinules, and it curves into a long tapering point. On the segment 
near the bases of these two setae are four spines which are much longer than in the 
other two species. The c, d and e setae are absent. The main setae are all thin 
walled and bare. The g seta is only a little longer than the others but appreciably 
broader. 

Synonomies. It is clear from Deevey's (1968) account that her material included 
specimens of this species; both from the size ranges and the number of spines in 
the armature of the e seta of the male antennule of her specimens described as C. 
procera Miiller. No other authors give sufficient information to determine the full 
range of this species. Miiller (i9o6a) also confused this species with C. procera. A 
re-examination of material from the Moroccan coast (Angel 1968) showed that the 
single adult female caught above the thermocline at 'Discovery' station 6183 haul 2 
(34i4'5'N, o803-o'W) belonged to this species. Recently more specimens were 
taken in the vicinity of I745'N, 2530'W (Angel unpublished). 



CONCHOECIA FROM THE NORTH ATLANTIC 



269 



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M. V. ANGEL 




FIG. 6. Conchoecia microprocera sp. nov., female. A, lateral view of carapace, B, ventral 
view of carapace; C, frontal organ and first antenna; D, second antenna; E, second 
antenna endopodite; F, male detail of the posterior dorsal corner of the carapace. 



Conchoecia tnacroprocera sp. nov. 

MALE. The holotype specimen mounted on slides in Euparal and stained with 
lignin pink has been deposited in the British Museum (Natural History) No. 1971.2.1.3. 
Collection data as for C. microprocera sp. nov. (see above). 



CONCHOECIA FROM THE NORTH ATLANTIC 




FIG. 7. Conchoecia macroprocera sp. nov., male. A, lateral view of carapace; B, ventral 
view of carapace; C, frontal organ and first antenna; D, first antenna detail of armature 
of the e, d and b setae; E, frontal organ capitulum; F, second antenna; G, left second 
antenna endopodite; H, right hook appendage; I, left hook appendage; J, sixth limb; 
K, penis. 



272 M. V. ANGEL 

Carapace (fig. 7 A). The range in size of 207 specimens was i- 10-1-22 mm with 
a mean of 1-153 0-021 mm. The relative height tended to be less and the breadth 
greater than for the other species. However, a distinguishing feature is that in all 
specimens the carapace breadth was greater than the height. The openings of the 
various glands are as described in the other species. There is no spine on the right 
valve at the posterior dorsal corner (fig. gF) (c.f. Miiller igo6a T. XIII fig. 37). 

Frontal organ (fig. 76). The shaft reaches level with the end of the limb of the 
first antenna. The capitulum is long and down turned. Two lateral thickening 
bars carry fine long hairs for the proximal third. The ventral edge has a small 
swelling on its median third, on the distal end of which is a more obvious group of 
long hairs. The dorsal edge is concave and bare. The end of the capitulum is 
rounded with a low subterminal knob (fig. 7E). One of Miiller's figures (igo6a 
T. XIII fig. 45) shows some resemblance to the frontal organ of this species. 

First antenna (fig. 76). The first segment is much shorter than the second; 
contrasting with the other two species. The ganglionic body, which lies in the region 
of the articulation between the two segments contains many yellow brown pigment 
corpuscles. The a seta lies back parallel with the limb reaching beyond the end 
of the second segment. The seta is significantly shorter than in C. microprocera, 
but not significantly so than in C. procera Miiller. The b and d setae are sub-equal 
and only carry six and three minute spinules respectively on a level with the distal 
end of the e seta armature. The e seta armature (fig. 7D) consists of 28-30 pairs 
of long slightly curved spines which increase in length slightly towards the base of the 
seta. Distal of the main armature are two pairs of distally pointing spinules. 

Second antenna (fig. 7F). The relative lengths of the protopodite and exopodite 
are similar to those in C. procera Miiller. The longest swimming seta is longer and 
more closely approaches the protopodite length than in the other species. The 
a seta on the endopodite (fig. 7G) is 2/3's the length of the b seta and has a few very 
fine spinules. The b seta carries more longer spinules. The c and d setae are nearly 
as long as the second segment. The e seta is minute. The g seta is only a little 
longer than the f seta. Both the g and f setae are bare and the g seta is slightly 
flattened. The h seta has a slight swelling near its base. The left hook appendage 
(fig. 7!) has a basal right angle and then curves gently a further 30 or so. Ter- 
minally it tapers asymmetrically to a blunt point. The right hook appendage 
(fig. 7H) curves through a total of 150. The end is swollen with subterminal 
ridging and ending in a blunt asymmetrical point. 

Mandible (fig. 8C). The toothed edge of the coxale (fig. 8 A) has ten low blunt 
teeth. The distal tooth list consists of two large teeth, the second of which is 
serrated, followed by about twelve small regularly sized teeth. The proximal list 
has twelve to fifteen teeth regularly diminishing in size across the list. 

The toothed edge of the basale (fig. 8B) consists of the usual two tubiform fol- 
lowed by six wedge teeth with a single broad outer tooth. The outer setae are short 
and fail to reach the level of the teeth. There are five rows of hairs arranged on the 
surface between the insertion of the outer setae and the two tubiform teeth. On 
the inner edge of the first endopodite segment is a single long seta. The second 
segment is bare on its outer edge. 



CONCHOECIA FROM THE NORTH ATLANTIC 




FIG. 8. Conchoecia macroprocera sp. nov. A, mandible tooth lists and toothed edge of 
the coxale ; B, mandible toothed edge of the basale ; C, mandible endopodite ; D, maxilla 
endopodite; E, labrum; F, fifth limb; G, caudal furca. 



274 M. V. ANGEL 

Maxilla (fig. 8D). There is a basal seta. On the endopodite the first segment 
carries six anterior, one lateral and three posterior setae. There are about six 
spinules on the distal edge of the segment near its articulation with the second 
segment. 

Labrum (fig. 8E). The hyaline membrane has a shallow rounded notch. 

Caudal f urea (fig. 8G). The first hook spines reach just short of the tips of the 
second pair. There is no unpaired seta dorsal of the hook spines. Between the 
two caudal lamellae is a covering of fine hairs. 

Penis (fig. 7K). The end of the organ is rounded and it contains four oblique 
muscles. 

FEMALE. The paratype mounted on slides in Euparal and stained with lignin 
pink has been deposited in the British Museum (Natural History) No. 1971.2.1.4. 
Collection data as for C. microprocera sp. nov. 

Carapace (figs gA, B). The range in length of 263 specimens was 1-26-1 -36 mm 
with a mean of 1-304 + 0-021 mm. The relative height of the carapace is noticeably 
less than for C. procera Miiller although its breadth is similar. The asymmetrical 
glands are positioned as in the other two species. There is no spine on the right 
valve at the posterior dorsal corner. 

Frontal organ (fig. gC). The total length of the organ is nearly three times the 
length of the limb of the first antenna. There is some demarcation into shaft 
and capitulum. Near the rounded tip there is a row of spines on the ventral surface. 

First antenna (fig. gC). The segmentation is indistinct and there is no additional 
armature. The a-d setae are more than half the length of the e seta. The e seta 
carries short spinules on its posterior edge and sparser long spinules on the distal 
half of its anterior edge. 

Second antenna (fig. gD). The ratio of the relative lengths of the segments of the 
limb are very similar to those for C. procera Miiller (Table i), although the longest 
swimming seta does tend to be longer. On the endopodite the a seta is bare and 
2/3's the length of the b seta. The b seta carries a few short spinules. On the 
segment near the bases of these setae are about eight short fine spinules. The 
second segment is almost bare with only about three or four hairs. All the main 
setae are bare and thin walled. The g seta is only a little longer than the others 
but is much broader. 

Synonomies. Miiller (igoSa) gave a size range for C. procera which suggests 
that his material included this species. Similarly Leveau's (1965) report of C. 
procera Miiller reaching a length of 1-30 mm may indicate its presence in the Mediter- 
ranean. Leveau reported two centres of abundance in C. procera Miiller at 200- 
100 m and 500-300 m. It seems possible that the deeper population may be this 
species. The species was absent from Deevey's (1968) material, but has turned up 
in very small numbers recently in samples taken near i745'N, 253o'W (Angel 
unpublished). Thus this species may not normally occur north of about i8N in 
the North Atlantic. 



CONCHOECIA FROM THE NORTH ATLANTIC 



275 



D 




FIG. 9. Conchoecia macroprocera sp. nov., female. A, lateral view of carapace; B, ventral 
view of carapace; C, frontal organ and first antenna; D, second antenna; E, second 
antenna endopodite; F, details of the posterior dorsal corner of the male carapace. 

Conchoecia vitjazi Rudjakov 1962 

MATERIAL. A single female was found in a 'Discovery' net (Currie & Foxton 
1957) haul from 'Discovery' station 4768, position 4O03'N, I957'W, depth 4750- 
4000 m, time 1402-1640 hrs, date 12 October 1961. Since no description is available 
in English, the specimen is described in full. 



276 M. V. ANGEL 

MALE. The male is unknown. 

FEMALE. Carapace. The length of the 'Discovery' specimen is 2-00 mm. 
Rudjakov's specimens ranged from 2-20-2-30 mm. The 'Discovery' specimen was 
too distorted to make accurate measurements of carapace height and breadth, 
but the height is about half the carapace length. The whole carapace is covered 
with a fine V-shaped sculpturing (fig. loA). The right asymmetrical gland opens 
on a small prominence (possibly an artefact) and the left asymmetrical gland about 
o-i mm from the posterior dorsal corner (fig. loA). The right valve carries a minute 
spine at the posterior dorsal corner. The outline of the carapace is very similar 
to Rudjakov's (1962) figure 2a. 

Frontal organ (fig. loB). The organ is much longer than the limb of the first 
antenna. There is no clear distinction into stalk and capitulum. Terminally the 
organ is down-turned and carries a few small spinules on the ventral corner (fig. loC). 

First antenna (fig. loB). The segmentation is indistinct and there is no addi- 
tional armature. The dorsal seta is absent. The e seta is over twice the length 
of the other setae and only carries a few fine spinules on the distal third of its trailing 
edge. These spinules were on the third quarter of the e seta in Rudjakov's speci- 
mens. 

Second antenna (fig. loE). The proportion of the protopodite : exopodite seg- 
ment i ; exopodite segments 2-8 is 6 : 3 : i in the 'Discovery' specimen and 4:2-8:1 
in Rudjakov's specimens. On the endopodite (fig. loF) the processus mamillaris 
is small and bluntly pointed. The a seta is bare and two thirds the length of the 
b seta. The b seta carries a group of long hairs near its base and very fine spinules 
for about half its length. The c, d and e setae are all absent. The g seta is as long 
as the protopodite, and is flattened terminally with fine spinules along the edge 
of the flattened part. The other setae are long subequal, thin-walled and without 
shafts. 

Mandible (fig. nB). The toothed edge of the basale is typical for the genus 
(fig. 1 1 A). The broad outer tooth is as broad as three of the other teeth. The 
outer setae project just beyond the level of the teeth. There are four groups of 
hairs near the bases of the tubiform teeth. Between the toothed edge and the 
articulation with the first endopodite segment is a patch of fine hairs. The inner 
edge of the first endopodite segment carries one long seta reaching well beyond the 
tips of the shorter of the terminal setae, and three minute setae arranged in a line 
up the segment. On the distal half of the outer surface of the outer segment is a 
patch of fine spinules. The toothed edge of the coxale has nine unusually elongate 
teeth (fig. loD). The distal list has two large teeth, the second of which is very 
serrated, followed by 17 small teeth. The proximal list has a large tooth followed 
by about six small teeth, another large tooth and a further 20 smaller irregular 
teeth. This is in substantial agreement with Rudjakov's (1962) type description. 
There is a seta on the distal outer edge of the first segment which Rudjakov noted 
as absent but was probably broken off in his specimens. 

Maxilla (fig. nC). The maxilla has a long basal segment. There are six anterior, 
one lateral and three posterior setae on the first segment. There are no spines 



CONCHOECIA FROM THE NORTH ATLANTIC 



277 



on the outer edge of the segment. The second endopodite segment is long and thin 
with long terminal hook setae. 





D 



FIG. 10. Conchoecia vitjazi Rudjakov, female. A, carapace right valve mounted flat; 
B, frontal organ and first antenna; C, detail of the tip of the frontal organ; D, mandible 
tooth lists and toothed edge of the coxale; E, second antenna; F, second antenna endopo- 
dite. 



2 7 8 



M. V. ANGEL 



Labrum. The hyaline membrane has a deep rounded notch. 

Caudal f urea. The first pair of hook spines just fail to reach level with the ends 
of the second pair. There is an unpaired seta dorsal of the hook spines. Between 
the lamellae is a covering of fine hairs. 




B 



FIG. n. Conchoecia vitjazi Rudjakov, female. A, mandible toothed edge of the basale; 
B, mandible endopodite; C, maxilla endopodite. 



DISCUSSION 



At 'Discovery' station 6665 C. procera Miiller was most abundant by day from 
50-300 m, C. microprocera sp. nov. was most abundant at 25 m and occurred in 
quite large numbers down to 100 m, C. macroprocera sp. nov. was most abundant 



CONCHOECIA FROM THE NORTH ATLANTIC 




FIG. 12. Conchoecia decipiens Miiller, male. A, lateral view of the carapace; B, ventral 
view of the carapace; C, frontal organ and first antenna; D, frontal organ capitulum; 
E, first antenna armature of the b, e and d setae; F, second antenna; G, left second 
antenna endopodite; H, right hook appendage; I, left hook appendage; J, sixth limb; 
K, penis; L, details of the tip of the penis. 



280 



M. V. ANGEL 



from 500-300 m and was absent from above 200 m. At night C. procera and C. 
microprocera showed reverse migrations out of the surface 50-75111, whereas C. 
macroprocera did not migrate. 




FIG. 13. Conchoecia decipiens Miiller. A, mandible tooth list and toothed edge of the 
coxale; B, mandible toothed edge of the basale; C, mandible endopodite; D, maxilla 
endopodite; E, labrum; F, fifth limb. 



CONCHOECIA FROM THE NORTH ATLANTIC 



281 



The other species which have been attributed to the 'procera' group are C. decipiens 
Miiller igoGa, C. brachyaskos Miiller I9o6a and C. vitjazi Rudjakov 1962. C. 
decipiens has only been reported from the Indian Ocean from between 23 N (Leveau 
1968) and 25 S (Miiller 1908). To show clearly the differences between C. decipiens 
Miiller and the other species, especially C. macroprocera sp. nov. drawings of speci- 
mens lent me by Mr J. George of the Cochin Indian Ocean Sorting Centre are in- 
cluded (figs 12, 13, 14). Table I also includes measurements of the specimens. 




FIG. 14. Conchoecia decipiens Miiller, female. A, lateral view of the carapace; B, ventral 
view of the carapace; C, frontal organ and first antenna; D, second antenna; E, second 
antenna endopodite; F, male, detail of the posterior dorsal corner of the carapace. 



282 M. V. ANGEL 

C. decipiens Miiller has a large spine and secondary spine on the right carapace 
valve at the posterior dorsal corner. In the males the following characters distin- 
guish C. decipiens from C. macroprocera: (i) The capitulum of the frontal organ is 
distinctive. (2) The first antenna is relatively longer in C. decipiens. (3) The 
first antenna e seta armature has no distally pointing spinules and the main spines 
decrease in size proximally. (4) On the second antenna the protopodite is longer 
and the shape of the right hook appendage is distinctive. (5) The endopodite of 
the mandible carries two setae on the inner face of the first segment. (6) The 
penis has five oblique muscles. 

In the females the following characters distinguish C. decipiens from C. macro- 
procera: (i) The outline of the carapace, (2) the shape of the frontal organ, (3) the 
endopodite of the second antenna has no spinules on the first segment near the bases 
of the a and b setae, the second segment is completely bare and the g seta is markedly 
flattened. 

Miiller (igo6a) included C. brachyaskos in the 'procera' group on the basis of the 
female frontal organ and the shortness of the sensory setae on the endopodite of 
the male second antenna. However, because of the setation of the male first 
antenna and its distinctive e seta armature (Miiller igo6a T. XIV fig. 12), it has not 
been included here. C. brachyaskos Miiller was present in the deep midwater samples 
from 'Discovery' station 6665. There are two size groups; the larger occurring 
only from below 1000 m. This large form is superficially very similar to the Ant- 
arctic form and so this species requires careful taxonomic investigation before its 
status can be certain. 

The discovery of a specimen attributable to C. vitjazi Rudjakov in the North 
Atlantic provides an interesting example of the faunistic relationships between the 
Pacific and the North Atlantic (c.f. Briggs 1970). This species, only recorded 
before from below 6000 m in the Kurile-Kamchatka Trench (463i'N, i5422'E, 
4348'N, i4955'E), was considered by Rudjakov (1962) to be endemic to that region. 
The 'Discovery' specimen does show some minor variations from the type description, 
but these are not considered great enough to attribute to it new specific rank. The 
male of the species is yet to be described and so the inclusion of the species in the 
'procera' group is yet to be confirmed. 



SUMMARY 

1. Conchoecia procera Miiller is shown to have been confused with two very 
closely related but distinct species in the past. 

2. C. procera sensu strictu is described and the two new species are described 
and named C. microprocera sp. nov. and C. macrorpocera sp. nov. 

3. An adult female ascribed to C. vitjazi Rudjakov 1962 is described from the 
North Atlantic ; a species previously thought to be endemic to the Kurile-Kamchatka 
Trench. 

4. The inclusion of C. brachyaskos Miiller in the 'procera' group is questioned. 



CONCHOECIA FROM THE NORTH ATLANTIC 283 

REFERENCES 

ANGEL, M. V. 1968. The thermocline as an ecological boundary. Sarsia 34 : 299-311. 

- i96ga. Planktonic ostracods from the Canary Island region; their depth distributions, 
diurnal migrations and community structure. /. mar. biol. Ass. U.K. 49 : 515-553. 

- ig69b. Repeated samples from a deep midwater planktonic ostracod community. /. 
exp. mar. biol. Ecol. 3 : 76-89. 

BRIGGS, J. C. 1970. A faunal history of the North Atlantic Ocean. Syst. Zoo/. 19 : 19-34. 
CLAUS, C. 1890. Die Gattungen und Arten der mediterranen und atlantischen Halocypriden 
nebst Bermerkungen liber die Organisation derselbe. Arb. Zoo/. Inst. Wien 9 : 1-34. 

- 1894. Die Halocypriden und ihre Entwicklungsstadien. Gesammelt 1890, 1891, 1892, 
1893. Denkschr. Akad. Wiss., Wien Math.-nat. Kl. 61 : i-io. 

CURRIE, R. I. & FOXTON, P. 1957- A new quantitative plankton net. J. mar. biol. Ass. 

U.K. 36 : 17-32. 
DEEVEY, GEORGIANA B. 1968. Pelagic ostracods of the Sargasso Sea off Bermuda. Bull. 

Peabody Mus. nat. Hist. 26 : 1-125. 
FOWLER, G. H. 1909. The Ostracoda. Biscayan Plankton, XII. Trans. Linn. Soc. Land., 

(2) (Zoo/.) 10 : 219-336. 
FOXTON, P. 1963. An automatic opening-closing device for large plankton nets and midwater 

trawls. /. mar. biol. Ass. U.K. 43 : 295-308. 

- 1969. SOND cruise 1965. Biological sampling methods and procedures. /. mar. 
biol. Ass. U.K. 49 : 603-620. 

GRANATA, L. & Di CAPORACCIO, L. 1949. Ostracodes marins recueillis pendant les croiseres 
du Prince Albert i er der Monaco. Res. Camp. sci. Monaco 109 : 1-51. 

HURE, J. 1955. Distribution annuelle verticale du zooplancton sur une station de 1'Adriatique 
meridionale. A eta Adriat. 7 : 3-69. 

- 1961. Distribution saisonniere et migration journaliere verticale du zooplancton dans la 
region profonde de 1'Adriatique. Acta Adriat. 9 : 3-59. 

ILES, E. J. 1953. A preliminary report on the Ostracoda of the Benguela Current. 'Dis- 
covery' Rep. 57 : 259-280. 

LEVEAU, M. 1965. Contribution a 1'etude des Ostracodes et Cladoceres du Golfe de Marseille. 
Rec. Trav. St. mar. Endoume 37 : 161-243. 

- 1967. Ostracodes pelagiques du sud-ouest de 1'Ocean indien (Regeon de Tulear). Rec. 
Trav. St. mar. Endoume, suppl. 6 : 63-70. 

- 1969. Ostracodes pelagiques recuellis lors de la 3 e campagne de 1'aviso 'Commandant 
Robert Giraud'. Rec. Trav. St. mar. Endoume, suppl. 8 : 127-280. 

MULLER, G. W. 1890. Ueber Halocypriden. Zoo/. Jb. Abt. System. 5 : 253-280. 

- 1894. Die Ostracoden des Golfes von Neapel und der Angrenzenden Meeres-abschnitte. 
Fauna Flora Golf. Neapel 21 : 29-154. 

I9o6a. Ostracoda. Wiss. Ergebn. Deutsch. Tiefsee-Exped. 8 : 29-154. 

I9o6b. Ostracoden der Siboga Expedition. Siboga Exped. 30 : 1-40. 

- 1908. Die Ostracoden der Deutschen Siidpolar Expedition 1901-1903. Sudpolar 
Exped. 10 : 51-182. 

1912. Ostracoden. Das Tierreich, 31 : 1-434. 

RUDJAKOV, J. A. 1962. Ostracoda Myodocopa of the family Halocypridae from the north- 
western Pacific. Trudy Inst. Okeanol. 58 : 172-201. (Translation available from N.I.O.) 

SCHWEIGER, L. 1912. Adriatische Cladoceren und Plankton-Ostracoden. Sber. Akad. Wiss. 
Wein, Math.-Nat. Kl. 121 : 239-272. 



Dr M. V. ANGEL 

NATIONAL INSTITUTE OF OCEANOGRAPHY 

WORMLEY, GODALMING, SURREY 




A LIST OF SUPPLEMENTS 
TO THE ZOOLOGICAL SERIES 

OF THE BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 



1. KAY, E. ALISON. Marine Molluscs in the Cuming Collection British Museum 
(Natural History) described by William Harper Pease. Pp. 96; 14 Plates. 
1965. (Out of Print.) 3.75. 

2. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and 
Valenciennes. Pp. 180; n Plates, 15 Text-figures. 1967. 4. 

3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure of Mineralogy 
at the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125; 29 Plates 
77 Text-figures. 1969. 4.50. 

4. HAYNES, J. R. Cardigan Bay recent Foraminifera (Cruises of the R.V. Antur) 
1962-1964. (In press.) 



Printed in England by Staples Printers Limited at their Kettering, Northants, establishment 




* 



I 

J MISCELLANIA 





BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 21 No. 8 

LONDON : 1972 



MISCELLANIA 




pp. 285-361 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 21 No. 8 

LONDON: 1972 



THE BULLETIN OF THE BRITISH MUSEUM 

(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 21, No. 8 of the Zoological 
series. The abbreviated titles of periodicals cited follow 
those of the World List of Scientific Periodicals. 



World List abbreviation 
Bull. Br. Mus. nat. Hist. (Zool.). 



Trustees of the British Museum (Natural History), 1972 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 4 February, 1972 Price 2.60 



CONTENTS 

Page 

Conchoecia pseudoparthenoda. By MARTIN VIVIAN ANGEL . . . 289 

Genus Lithobius. By EDWARD HENRY EASON 297 

The Chestnut-shouldered Wren. By COLIN JAMES OLIVER HARRISON . 313 

Lironeca (Isopoda; Cymothoidae). By ROGER JOHN LINCOLN . . . 329 

Eunice manihine (Polychaeta; Eunicidae). By MICHAEL ROY LONGBOTTOM 339 

The Gangetic Dolphin Platanista gangetica. By GEORGE PILLERI . . 345 

A new Genus and Species of Sudan Leech. By LAURENCE REGINALD 

RICHARDSON. .......... 349 

Genera Phoxinellus, Pseudophoxinus and Paraphoxinus. By ETHELWYNN 

TREWAVAS 359 



CONCHOECIA PSEUDOPARTHENODA (NOV. SP) 

A NEW HALOCYPRID OSTRACOD FOR THE 

TROPICAL NORTH ATLANTIC 

By M. V. ANGEL 

INTRODUCTION 

IN the Spring of 1968 a series of horizontal tows were made by R.R.S. Discovery at 
station 6665 to study the vertical distribution of planktonic animals at 10 16' N, 
19 47' W. A new Conchoecia species belonging to the magna group and very 
closely related to C. parthenoda Miiller 1906 occurred in the near surface hauls. 
The species is described here and named Conchoecia pseudoparthenoda. 

Conchoecia pseudoparthenoda n. sp. 

The type specimens were from haul 36 at station 6665 from a depth of 25 m at 
1637-1715 hrs on 26 February 1968, position 10 16' N, I947'W. They are 
deposited mounted on slides in the British Museum (Natural History), male No. 
1971.2.1.5, female No. 1971.2.1.6. 

MALE. The range in carapace length of 120 specimens was 1-56-1-72 mm with 
a mean of 1-637 '37 mm. The outline of the carapace (Fig. lA) is similar to 
that of C. parthenoda Miiller (Deevey 1968, Angel I969a). The position of the left 
asymmetrical gland is quite distinctive; in the type specimen it opens 0-48 mm 
anterior of the posterior carapace hinge. The right asymmetrical gland opens at 
the usual position on the posterior ventral corner. Fine spines occur along the 
dorsal surface of the carapace in the region of the hinge line, and on the edges of the 
valves from below the rostral incisure to about the midpoint of the ventral side. 

Frontal organ. The shaft extends to level with the end of the limb of the first 
antenna (Fig. iB). The capitulum is down-turned, its posterior edge is almost 
straight and carries spines down the proximal two thirds (Fig. iC). The anterior 
edge is slightly concave so that the capitulum is narrowest in its middle region, and 
carries a few spines near its base. The end of the capitulum is rounded. 

First Antenna. The two basal segments are sub-equal and bare (Fig. iB). The 
a seta lies back parallel with the limb reaching level with or just short of the joint 
between the two basal segments. The c seta is short. The b seta carries nine fine 
spines level with the distal end of the armature on the e seta (Fig. iD), whereas the 
d seta carries about 27 fine spines. The b seta is only slightly shorter than the d 
seta, which is only just shorter than the e seta. The e seta armature consists of 9 
to 10 paired spines followed by 22-26 alternating spines, with a total range of 40-44 
spines (Fig. iD). The e seta also has a few scattered spines on its anterior leading 
edge close to its base. 

Second Antenna. The protopodite is more than half the length of the carapace 

Bull. Br. Mus. nat. Hist. (Zool.) 21, 8 



2 go 



M. V. ANGEL 



and three times the first exopodite segment (Fig. lE). All the exopodite segments 
are bare. The longest swimming seta is two fifths the carapace length. On the 
endopodite, the a seta is bare and curves back behind the b seta (Fig. iG). The b 





FIG. i. Conchoecia pseudoparthenoda male. A. Outline of carapace. B. Frontal organ 
and first antenna, c. Capitulum of frontal organ. D. Detail of armature of the anten- 
nular e, b and d setae. E. Second antenna. F. Copulatory organ. G. Endopodite of 
the left second antenna. H. Endopodite of the right second antenna. 



CONCHOECIA PSEUDOPARTHENODA (NOV. SP.) 291 

seta carries three long hairs near its base and fine spines distally. The processus 
mamillaris is bluntly pointed. The c and d setae are sub-equal and rather short. 
The e seta is minute. The g seta is almost as long as the protopodite and the f seta 
is similar in length to the longest swimming seta. The h, i, and j setae are half the 
length of the f seta and about two-fifths the length of the g seta. They have weakly 
developed shafts. The right hook appendage is right angled near its base, curves 
through a further 90 half way along its length ending in a point with subterminal 
ridging (Fig. iH). The left hook appendage is bent through less than a right angle 
near its base and is then straight (Fig. iG). Towards its end it narrows and ter- 
minates in a curved point with subterminal ridging. 

Mandible. The basal segment is long (Fig. zC). The first exopodite segment 
carries two setae on its inner edge, one twice the length of the other. The toothed 
edge of the pars incisa has two spine teeth followed by six broad finely serrated 
teeth, and a single inner broad tooth (Fig. 26). Hairs are inserted near the bases of 
the spine teeth and the outer setae. These setae project only just beyond the level 
of the teeth. The coxale toothed edge has ten bluntly pointed teeth (Fig. 2 A). 
The distal tooth list has two large teeth ; the second is serrated and followed by 13 
small blunt teeth. The proximal list has about 14 irregularly sized teeth. 

Maxilla. The basal segment carries a seta (Fig. 2D). The first endopodite 
segment has six anterior, one lateral and three posterior setae. There is a group of 
about six short spines at the end of the segment. 

Labrum. This is the usual shape for the magna group (Fig. 2E). 

Caudal Furca. The furca has eight pairs of hook spines (Fig. 2F). The first pair 
does not reach the level of the ends of the second pair. There is no unpaired post- 
erior seta. 

Copulatory organ. This has eight oblique muscles (Fig. iF). The terminal edge 
which curves in and over the intromittent spine is serrated. 

FEMALE. The range in length of 222 specimens was i -72-1 -90 mm with a mean 
of 1-841 0-038 mm. As in the male the left asymmetrical gland opens 0-48 mm 
anterior of the posterior hinge, and the right gland on the posterior ventral corner 
(Fig. 3 A). Fine spines occur on the dorsal surface in the vicinity of the hinge line 
and round the ventral edges of the valves in the anterior half. In many specimens 
there were concentric striations clearly visible over the whole of the carapace. 

Frontal organ. It is straight with no separation into shaft and capitulum (Fig. 
36). It terminates in a long downturned point. There are a few spines on its 
ventral edge (Fig. 36). 

First Antenna. The segmentation of the limb is indistinct. The long spinous 
dorsal seta reaches almost level with the tip of the frontal organ (Fig. 36, C). There 
is a group of spinules close to the insertion of the e seta. The e seta is twice the 
length of the limb and carries spines on its trailing posterior edge distal of the ends 
of the other setae. The other four setae are a third the length of the e seta. 

Second Antenna. The protopodite is just less than half the carapace length and 
nearly three times the length of the first exopodite segment (Fig. 3D). All the 
exopodite segments are bare. The longest swimming seta is three quarters the 
length of the protopodite. On the endopodite the a seta is half the length of the 



292 



M. V. ANGEL 



b seta, and both are finely spinous (Fig. 3E). The processus mamillaris has a small 
pointed tubercle at its tip. The g seta is just over half the length of the protopodite. 
It is unflattened and carries fine spinules near its tip. The f seta is only a little 
longer than the h, i and j setae which are two thirds the length of the g seta. 





D 




B 005 




FIG. 2. Conchoecia pseudoparthenoda. A. Toothed edge of coxale and tooth lists of the 
mandible. B. Toothed edge of the mandibular basale. c. Endopodite of the mandible. 
D. Maxilla endopodite. E. Labrum. F. Caudal furca. 



CONCHOECIA PSEUDOPARTHENODA (NOV. SP.) 



293 



JUVENILES. Two juvenile stages were recognised from the hauls by the position 
of the left asymmetrical glands. The smallest group consisting of 106 specimens 
ranged in size from 0-86-0-96 mm with a mean of 0-903 + 0-022 mm. The final 
stage juveniles ranged in size from 1-22-1-34 mm with a mean of 1-286 + 0-026 mm 
for 163 specimens. 

COMPARISON BETWEEN C. pseudoparthenoda AND C. parthenoda. In the females, 
C. pseudoparthenoda often has more distinctive sculpturing and the posterior ventral 
region of the carapace is more strongly developed. In both sexes the left asym- 








FIG. 3. Conchoecia pseudoparthenoda female. A. Outline of carapace. B. Frontal organ 
and first antenna, c. Details of frontal organ and the end of the first antenna. D. Second 
antenna. E. Endopodite of the second antenna. 



294 M - v - ANGEL 

metrical gland is 0-48 mm anterior of the posterior hinge, and this is diagnostic of 
the species; in C. parthenoda the gland opens 0-24 mm anterior of the posterior 
hinge. The carapace breadth is less in C. pseudoparthenoda. The shapes of the 
frontal organs of the females of the two species are quite distinct. The dorsal seta 
of the first antenna is shorter in the female of C. parthenoda and this species has no 
spinules at the insertion of the first antennal e seta, also the e seta is relatively 
longer. Similarly the exopodite segments and the endopodite setae of the second 
antennae of the females are all relatively longer in C. parthenoda. A comparison 
of the lengths of the various setae and other meristic characters of the two species 
are shown in Table i. 

In the males C. parthenoda has a longer b seta on the first antenna and a shorter 
c seta ; the b and d setae carry more spines. The antennular e seta armature usually 
consists in both species of nine pairs of distal spines. In C. parthenoda the paired 
spines are followed by about 18 alternating spines; giving a total range of 35-38 
spines. In C. pseudoparthenoda the paired spines are followed by about 24 alter- 
nating spines ; giving a total range of 40-44 spines. 

On the mandibles the proximal tooth lists are quite distinct ; C . parthenoda having 
three large teeth and 16 smaller teeth, and C. pseudoparthenoda 14 irregular teeth. 
On the caudal furca C. parthenoda has an unpaired dorsal seta but C. pseudopar- 
thenoda has none. The live colouration of the two species is similar, for the most 
part they are transparent and colourless or very pale red. The anterior ventral 
edges of the carapace valves are yellow, and the pharyngeal region is yellow or 
orange. 

Bioluminescence has not been observed in either of the two species. 

DISCUSSION 

At station 6665 C. pseudoparthenoda predominated over C. parthenoda at 25 m, 
but C. parthenoda was the more numerous in the 50 m and 75 m hauls. Both 
species were virtually restricted to the surface 100 m. C. pseudoparthenoda was 
absent from the plankton off Fuerteventura (Canary Islands) and the Moroccan 
coast (Angel 1968, 1969^, but Deevey (personal communication) has found what is 
probably this species to be a rare member of the plankton off Bermuda. C. par- 
thenoda was common in all these areas. 

Poulsen (1969) has recently described a male specimen from the Congo region 
with the gland openings of the left asymmetrical gland 0-38 mm from the posterior 
dorsal hinge (see his figure 10 and table p. 154). The females, however, had gland 
openings 0-27 mm from the hinge. The carapace lengths of all his specimens were 
slightly larger than described here for C. pseudoparthenoda, and appreciably longer 
than for C. parthenoda. His description of the male antennular e seta armature 
makes no mention of the distal spines being paired, and the total counts are con- 
siderably lower than for either C. parthenoda or C. pseudoparthenoda. It seems 
unlikely from his description that Poulsen's specimens belonged to either of these 
species. 

C. pseudoparthenoda appears to have an equatorial distribution in the Atlantic 
and may prove to be endemic to this region. 



CONCHOECIA PSEUDOPARTHENODA (NOV. SP.) 



295 



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REFERENCES 

ANGEL, M. V. 1968. The thermocline as an ecological boundary. Sarsia 34 : 299-312. 

- igGga. The redescription of three halocyprid ostracods, Conchoecia hyalophyllum Claus, 
C. magna Claus and C. parthenoda Mullet from the North Atlantic. Crustaceana 17, pp. 

45-63- 

- 19695. Planktonic ostracods from the Canary Island region: their depth distributions, 
diurnal migrations and community organization. /. mar. biol. Ass. U.K. 49 : 515-553. 

DEEVEY, Georgiana. 1968. Pelagic ostracods of the Sargasso Sea off Bermuda. Bull. 

Peabody Mus. nat. Hist. 26 : 1-125. 

MULLER, G. W. 1906. Ostracoda. Wiss. Ergebn. Deutsch. Tiefsee-Exped. 8 (2), pp. 29-154. 
POULSEN, E. M. 1969. Ostracoda-Myodocopa from the Eastern Tropical Atlantic. Vidensk. 

Meddr. dansk naturh. Foren. 132 : 129-197. 



Dr. M. V. ANGEL 

NATIONAL INSTITUTE OF OCEANOGRAPHY 

WORMLEY, 

GODALMING, SURREY 



THE TYPE SPECIMENS AND IDENTITY OF THE 

SPECIES DESCRIBED IN THE GENUS 

LITHOBIUS BY GEORGE NEWPORT IN 1844, 

1845 AND 1849 (CHILOPODA, LITHOBIOMORPHA) 

By E. H. EASON 

INTRODUCTION 

GEORGE NEWPORT described eighteen nominal species under Lithobius one of which, 
L. emarginatus, he later removed to the genus Henicops. Pocock (1890, iSgia, 
i8gib, 1901) examined the type specimens of at least eight of these species but he 
only identified five, one of them incorrectly. Of the remainder, some have been 
identified more or less definitely by various authors from their original descriptions 
but the identity of the others has only been tentatively suggested or has, hitherto, 
been quite unknown. The type specimens of sixteen of Newport's species are 
preserved either in the British Museum (Natural History), the Hope Department of 
Zoology, Oxford or the Museum National d'Histoire Naturelle, Paris. They have 
all been re-examined for the purpose of the present study and their identity is either 
confirmed or established for the first time. It is not always possible to tell whether 
Newport had one or several specimens before him when he wrote his descriptions and 
he made no formal designation of type specimens, but where only a single specimen 
is available it is regarded as the holotype: otherwise a lectotype is selected where 
necessary. An attempt is also made to determine the identity of the species for 
which type specimens have not been found. 

Newport's 1844-45 paper was divided between parts 3 and 4 of volume 19 of the 
Transactions of the Linnean Society of London. The section in part 3, which includes 
a figure with the caption 'Lateral view of the head of Lithobius americanus, Newp.' 
(Tab. 33), was published in November 1844, but the section in part 4, which includes 
the written descriptions of this and eight other new species of Lithobius, was not 
published until November 1845 (Raphael, 1970). The species described in this paper 
are usually dated 1844, but clearly all except L. americanus should be dated 1845. 

Conclusions as to the status and present classification of all the nominal species 
described by Newport in the genus Lithobius are summarized in Table i. 

i. Lithobius hardwickei Newport 
Fig. i 

Lithobius Hardwickei Newport, 1844, p. 96; 1845 (1844-45), p. 366 

TYPE LOCALITY. Singapore. 

TYPE SPECIMEN. Holotype : a male pseudomaturus of L.forficatus (Linn.) 16 -5 mm 
long, dried and pinned, labelled "L. Hardwickei Newp." in Newport's hand and 

Bull. Br. Mus. nat. Hist. (Zool.) 21, 8 



298 E. H. EASON 

"Hardwicke Bequest, Singapore" on a separate ticket. British Museum (Natural 
History). 

REMARKS. As Newport stated, this specimen is smaller than an adult of L. 
forficatus and of a paler colour than usual : it has only 39 antennal articles (Newport 
gave 41) but the prosternum is exactly as described by Newport with 5+8 teeth 

(Fig. i). 

According to Haase (1887), Pocock examined the specimen and found it to have 
posterior projections on T. 9, n and 13 but came to no conclusion as to its identity. 
Verhoeff (1937) merely noted that Newport's description of L. hardwickei agreed 
with none of the species of Lithobius he found in the Malay peninsula. Wang and 
Tang (1965), in the most recently published list of Chilopoda from Singapore, made 
no mention of the species and there is little doubt that L. forficatus was introduced 
to Singapore but never became established. 

2. Lithobius leachi Newport 

Lithobius forficatus : Leach, 1814, p. 408 

Lithobius Leachii Newport, 1844, p. 96; 1844 (1844-45), Tab. 33, fig. 30 

Lithobius sp. Newport, 1844 (1844-45), Tab. 33, fig. 31 

Lithobius Leachii: Newport, 1845 (1844-45), p. 368 

TYPE LOCALITY. Europe. 

TYPE SPECIMEN. Holotype: a female of L. forficatus (Linn.) 20 mm long, dried 
and gummed to a card over the whole of its ventral aspect, labelled "L. Leachii 
Newp." in Newport's hand. British Museum (Natural History). 

REMARKS. Newport examined the original Linnean specimen of L. forficatus and 
finding it to differ slightly in the form of the prosternum from Leach's specimen of 
this species, referred the latter to a new species, L. leachi. Synonymy of L. leachi 
with L. forficatus was tentatively suggested by Meinert (1868) and has never been 
disputed. Although the ventral aspect of the holotype is obscured owing to its 
being gummed to a card, it is undoubtedly the specimen which Newport examined 
and which Leach had, quite correctly, identified as L. forficatus. 

3. Lithobius pilicornis Newport 

Lithobius pilicornis Newport, 1844, p. 96 

Lithobius sp. Newport, 1844 (1844-45), Tab. 33, fig. 34 

Lithobius pilicornis : Newport, 1845 (1844-45), p. 369 

TYPE LOCALITY. England. 

TYPE SPECIMEN. Holotype: a male 26 mm long, dried and pinned, labelled "L. 
pilicornis Newp." in Newport's hand. British Museum (Natural History). 

REMARKS. This specimen was described by Pocock (iSgia) who gave a full 
account of the species. 

Gervais, in his list of published descriptions of species of Myriapoda (Walckenaer 
and Gervais, 1847), mentioned Lithobius pulchricornis and L. pilicornis as having 



GENUS LITHOBIUS 



299 



been described by Newport in 1844 and 1845 respectively. It seems that pulchri- 
cornis attaches to Newport's first (1844) brief description of L. pilicornis. The 
name does not appear in any of Newport's works and must have arisen from some 
carelessly written transcript of his earlier paper. 



\ 







FIGS 1-5. i, Lithobius hardwickei. Dental margin of prosternum of holotype, ventral. 
3, Lithobius americanus. Dental margin of prosternum of holotype, ventral. 2, Litho- 
bius brevicornis. Dental margin of prosternum of holotype, ventral. 4, Lithobius 
elongatus. gth to i4th tergites of lectotype, dorsal. 5, Lithobius monilicornis. gth to 
1 4th tergites of male syntype, dorsal. 



3 oo E. H. EASON 

4. Lithobius sloanei Newport 

Lithobius Sloanei Newport, 1844, p. 96; 1845 (1844-45), p. 369 
TYPE LOCALITY. Unknown. 

TYPE SPECIMEN. Holotype: a bleached and distorted female of L. pilicornis 
about 30 mm long, dried and pinned, labelled "L. Sloanei Newp." in Newport's 
hand and "Sir H. Sloane's coll. ?4i67" on a separate ticket. British Museum 
(Natural History). 

REMARKS. This specimen was described and discussed by Pocock (iSgia) who 
identified it correctly and noted that the number 4167 corresponds to the following 
entry in Sir Hans Sloane's catalogue "a middling good sized brown Scolopendra". 

5. Lithobius castaneus Newport 

Lithobius castaneus Newport, 1844, p. 96; 1845 (1844-45), p. 370 

TYPE LOCALITY. Sicily. 

TYPE SPECIMEN. Holotype: a female 20 mm long, dried and pinned, labelled "L. 
castaneus Newp." in Newport's hand. British Museum (Natural History). 

REMARKS. This specimen was examined by Pocock (1890) who recognized its 
identity with L. eximius Meinert, 1872. He noted correctly that it only has 26 
antennal articles (Newport gave 41). It agrees with Brolemann's (1930) description 
of L. castaneus except for the presence of the spine 15 VaT which was recorded by 
Meinert for L. eximius, and the extension of the distinctive sculpturing of the large 
tergites, rather faintly, on to T. 8, 10 and 12 : Brolemann described this last feature 
on T. 1.3 and 5 only. Other characters possessed by the holotype, all in agreement 
with Brolemann's description, are 6, 7, 7, 7 oblong coxal pores, lateral spines on the 
i4th and I5th coxae, and a simple claw on the gonopod with only a feeble lobe at the 
base of its external ridge and no denticles. These features place it in the subspecies 
L. castaneus buchnerorum described from Ischia by Verhoeff (1942) who assumed that 
the typical form of the species has a dentate female genital claw (Verhoeff, 1934). 
However, as Matic (1961) pointed out, many of the subspecies of L. castaneus 
described by Verhoeff were based on unstable characters and are probably without 
validity : should it be found that a geographical race with a dentate genital claw can 
be defined it can not be regarded as belonging to the nominate subspecies. 

6. Lithobius emarginatus Newport 

Lithobius emarginatus Newport, 1844, p. 96 

Henicops emarginatus: Newport, 1845 (1844-45), p. 372 

TYPE LOCALITY. New Zealand. 

TYPE SPECIMEN. Holotype: a very defective female 8-5 mm long, labelled "H. 
emarginatus Newp." and "in the ground Capt. Ross" in Newport's hand: the speci- 
men has been pinned through T. 7 damaging this and the adjacent tergites, but the 
pin has been withdrawn and the specimen is gummed to the apex of a triangular 
piece of card. British Museum (Natural History). 



GENUS LITHOBIUS 



301 



REMARKS. Pocock examined this specimen, redescribed the genera Henicops 
Newport and Lamyctes Meinert, and removed emarginatus to the latter (Pocock, 
1901). Archey (1937) also saw the specimen which he described as being 'much 
shrivelled and somewhat mutilated'. He noted however that the 2+2 prosternal 
teeth were discernible in the type and it is confirmed that in this as in all other 
respects the specimen is identical with the common New Zealand species Archey 
described under Lamyctes emarginatus. 

7. Lithobius rubriceps Newport 

Lithobius rubriceps Newport, 1845 (1844-45), p. 364 
TYPE LOCALITY. Southern Spain. 

REMARKS. Although Newport deposited his material belonging to this species in 
the British Museum, no type specimen has been found. It seems significant that 
Pocock, who examined and reported on most of Newport's type specimens of 
Lithobius in the British Museum, made no mention of L. rubriceps in any of his 
writings: this suggests that the specimen had already been lost or badly damaged 
before 1890 when Pocock made his first observations. 

However, the space in the cabinet containing Newport's specimens and apparently 
allotted to L. rubriceps is occupied by four examples of the common Iberian species 
known as L. insignis Meinert, labelled "Lithobius rubriceps Newport, Lisbon" and 
"96.3.8.103-106, Pascoe". These specimens were collected by the entomologist 
F. P. Pascoe and presented to the Museum by Miss Pascoe. They are accompanied 
by two further specimens of L. insignis labelled "Cintra 25.11.96". Newport's 
description is unmistakable and there is no doubt that these specimens were correctly 
identified, probably by Pocock, and that L. rubriceps is the senior synonym of 
L. insignis. 

Meinert (1872) described L. insignis and L. gracilipes as new species from Spain, 
suggesting L. rubriceps as a possible synonym of the latter. This choice on the part 
of Meinert is difficult to understand because Newport's description of rubriceps is 
much closer to insignis than to gracilipes. However, gracilipes only differs from 
insignis in being smaller with fewer coxal pores and may well prove to be another 
synonym of L. rubriceps. 

The species was fully described by Machado (1952). 

8. Lithobius fasciatus Newport 

Lithobius fasciatus Newport, 1845 (1844-45), p. 365 
TYPE LOCALITY. Florence and Naples. 

TYPE SPECIMENS. Lectotype: a specimen labelled "Lithobius fasciatus Newp." in 
Newport's hand. Paralectotypes: two unlabelled specimens accompanying the 
lectotype. Hope Department of Zoology. 

REMARKS. These three specimens have been fully described in an earlier paper 
(Eason, 1970). They were examined by Pocock (1890) who was mistaken in 
believing them to be identical with L. grossipes C. L. Koch. 



302 E. H. EASON 

9. Lithobius multidentatus Newport 

Lithobius multidentatus Newport, 1845 (1844-45), p. 365 
TYPE LOCALITY. New York. 

TYPE SPECIMEN. Holotype: a female 22 mm long labelled as the holotype of 
Lithobius multidentatus Newport by Dr. R. E. Crabill who removed it from the 
cabinet containing Newport's dried specimens in 1960, relaxed it in trisodium 
phosphate and placed it in spirit. British Museum (Natural History). 

REMARKS. Wood (1865) recognized the identity of L. multidentatus with the 
species he had himself described under Bothropolys nobilis (Wood, 1863), apparently 
basing his conclusion on Newport's very brief description. In fact, the holotype has 
9 + 9 prosternal teeth and not 8 + 8 as Newport stated. The specimen does, 
however, agree in all respects with Chamberlin's (1925) detailed description of 
Bothropolys rmdtidentatus except for the presence of a small extra medial spur on 
the left gonopod, in addition to the usual two. 

It is clear from Dr. Crabill's labelling of the holotype that he was satisfied as to its 
identity with the common North American species generally known as B. multi- 
dentatus. 

10. Lithobius americanus Newport 

Fig. 3 
Lithobius americanus Newport, 1844 (1844-45), Tab. 33, fig. 29; 1845 (1844-45), p. 365 

TYPE LOCALITY. North America. 

TYPE SPECIMEN. Holotype: a rather defective male of L. forficatus (Linn.) 24 mm 
long, dried and pinned, labelled "Lithobius Americanus Newp. N.S." in Newport's 
hand. Hope Department of Zoology. 

REMARKS. Newport distinguished this specimen from L. forficatus by its larger 
size and the form of the prosternum. The teeth of the latter are, indeed, irregularly 
spaced (Fig. 3) as in so many examples of L. forficatus. 

Synonymy of L. americanus with L. forficatus was first proposed by Stuxberg 
(1871) and has never been disputed. 

ii. Lithobius planus Newport 

Lithobius sp. Newport, 1844 (1844-45), Tab. 33, fig. 32 
Lithobius planus Newport, 1845 (1844-45), p. 366 

TYPE LOCALITY. North America. 

TYPE SPECIMEN. Holotype: a rather defective male of Bothropolys multidentatus 
18 mm long, dried and pinned, labelled "Lithobius planus Newp." in Newport's 
hand. Hope Department of Zoology. 

REMARKS. Newport made no use of the arrangement of the coxal pores in his 
system and so failed to detect the affinity between L. planus, which he believed to be 
close to L. variegatus Leach, and L. multidentatus. Further, the holotype of planus 



GENUS LITHOBIUS 303 

has 7 -f 7 prosternal teeth whereas multidentatus was described as having 8 +8 which 
is the more usual number in this species. Wood (1863, 1865) mentioned L. planus 
but merely reiterated Newport's description and made no suggestion as to its 
identity with any North American species known to him. 

12. Lithobius argus Newport 

Lithobius Argus Newport, 1845 (1844-45), p. 369 
TYPE LOCALITY. Wellington, New Zealand. 

TYPE SPECIMENS. Syntypes: a female pseudomaturus 18 mm long and a male 
praematurus 12-5 mm long, both of L. forficatus (Linn.), dried and pinned, labelled 
"Lithobius Zelandicus Newp." in Newport's hand. Hope Department of Zoology. 

REMARKS. Newport distinguished this form from L. forficatus by its smaller 
size ; only the female answers closely to his description. 

Pocock (iSgib) examined these specimens, noted that they were labelled "Litho- 
bius Zelandicus" , and confirmed that they belonged to Lithobius s.s. but came to no 
conclusion as to their exact identity. It seems that Newport first named them 
Zelandicus and then changed the name to Argus when writing his paper. 

This species has never been rediscovered in New Zealand but Archey (1937), in 
his most recently published account of the Chilopoda of that country, never ques- 
tioned its validity and regarded Newport's record as evidence of the occurrence of 
an indigenous species of Lithobius in a country where the Chilopod fauna consists 
otherwise almost entirely of genera confined to the southern hemisphere. There is 
no doubt that L. forficatus was introduced to New Zealand but lack of any subsequent 
records suggests that it has never become established. 

13. Lithobius brevicornis Newport 
Fig. 2 

Lithobius brevicornis Newport, 1845 (1844-45), p. 370 

TYPE LOCALITY. Naples. 

TYPE SPECIMEN. Holotype : a male pseudomaturus of L. forficatus (Linn.) 17 mm 
long, dried and pinned, labelled "Lithobius brevicornis Newp." in Newport's hand. 
Hope Department of Zoology. 

REMARKS. This specimen has 43 antennal articles, 15 ocelli on each side and 
4 +6 prosternal teeth (Fig. 2) , whereas Newport gave 41 articles, 20 ocelli and 6 -f-6 
teeth. There is, however, no suggestion in the description that it was based on more 
than one specimen and this description seems, therefore, to have been inaccurate. 
Newport equated L. brevicornis with L. vesuvianus Costa, but no account of the latter 
appears ever to have been published. 

Fanzago's (1874) and Fedrizzi's (1877) accounts of L. brevicornis are mere reitera- 
tion of the original description and Fedrizzi's mention of the antennae as having 14 
articles was obviously due to a misprint. Although L. brevicornis figures in a 



304 E. H. EASON 

number of more recent Italian faunal lists it has never been redescribed and no 
suggestion has ever been made as to its identity. 

14. Lithobius melanops Newport 

Lithobius melanops Newport, 1845 (1844-45), p. 371 
TYPE LOCALITY. Sandwich, England. 

TYPE SPECIMEN. Holotype : a very defective female 13 mm long, dried and pinned, 
labelled "Lithobius melanops Newp. (Kent 44.41)" in Newport's hand. British 
Museum (Natural History). 

REMARKS. Pocock (1890) examined this specimen and concluded that it was 
conspecific with a common European species widely referred to as L. glabratus C. L. 
Koch 1847. Although the antennae and most of the legs are missing there is no 
doubt that Pocock was correct. 

The species was described fully by Brolemann (1930). 

15. Lithobius platypus Newport 

Lithobius platypus Newport, 1845 (1844-45), p. 371 

TYPE LOCALITY. Egypt. 

REMARKS. On the single plate devoted to Myriapoda and Hexapoda Aptera in 
the Natural History portion of Savigny's "Description de 1'Egypte", figure 3 
represents a small lithobiomorph centipede, 11-5 mm long with short antennae each 
of 20 articles, i +2, i ocelli on each side, no posterior tergal projections and with 
both i4th and I5th legs thickened. Newport based his description of L. platypus 
on this figure which, with the thickened posterior legs, was unlike any of the larger 
species of Lithobius with which he was familiar. Although, like all Savigny's 
drawings, this figure is beautifully executed it is inaccurate in that a short tergite is 
shown immediately behind T. 7; such a tergite is never found in Lithobius or any 
related genus. On the other hand, apart from this spurious tergite, the figure is a 
very fair representation of L. vosseleri Verhoeff, originally described by Verhoeff 
(1901) from Cyprus and recorded by Silvestri (1929) from a number of localities in 
North Africa including Alexandria (Egypt). Silvestri referred his specimens to a 
new variety, propitia, on the basis of some quite trivial characters and suggested the 
possibility of their being identical with L. platypus, the only other species of Lithobius 
ever recorded from Egypt. 

Savigny's sight failed after he had completed his drawings but before he had 
finished his manuscript (Sherborn, 1897), so that no names, descriptions or exact 
localities attaching to his specimens of Myriapoda were ever published. Gervais 
(1837) commented on Savigny's figure of a lithobiid, but he did not give it a Latin 
binomen. Newport's description of L. platypus may, therefore, be the earliest 
published description of L. vosseleri, but there is no certainty of this and L. platypus 
should be rejected as a nomen dubium. 



GENUS LITHOBIUS 305 

16. Lithobius elongatus Newport 
Fig. 4 

Lithobius elongatus Newport in Lucas, 1849, p. 383, PI. 3, figs. 2, 2a, 2b, ac and 2d 

TYPE LOCALITY. Lac Tonga, Lac Houbeira and La Calle, Algeria. 

TYPE SPECIMENS. Lectotype: a male 24 mm long, preserved in spirit, labelled 
"Lithobius elongatus Newport, Algerie Lucas" is here formally designated as the 
lectotype. Paralectotypes : a distorted male about 28 mm long and two males 19 mm 
long accompanying the lectotype but now placed in a separate tube. Museum 
National d'Histoire Naturelle. 

REMARKS. Newport's description of L. elongatus is inadequate by modern 
standards but this species was more fully described by Verhoeff (1891) under 
Lithobius (Polybothrus) koenigi and by Silvestri (1896), in both instances from 
Tunisian specimens agreeing in all essentials with the lectotype. Later Silvestri 
(1897) described more specimens from Sicily and proposed elongatus and koenigi, 
together with L. impressus C. L. Koch and L. monilicornis Newport, as junior 
synonyms of L. nudicornis Gervais. L. nudicornis, however, is based on an 
extremely scanty description of a specimen from Sicily (Gervais, 1837) an d the name 
has been rejected by most authors in favour of elongatus. Whether impressus and 
monilicornis, which are usually accepted as synonyms of elongatus, should in fact be 
regarded as such is open to question and will be discussed under the next species. 
Owing to the uncertainty as to the relative status, not only of elongatus and 
impressus (=monilicornis) but of the numerous subspecies which have been des- 
cribed from time to time, the lectotype of L. elongatus is described below. 

DESCRIPTION OF LECTOTYPE. Size: 24 mm long, 2.75 mm broad at T. 10, i5th 
legs 8 mm long. Colour: bleached with little pigmentation. Antennae: 10 mm 
long with 44 articles. Ocelli: apparently 1+4, 4, 3 but difficult to see owing to 
bleaching. Prosternum : with 5 +5 faintly pigmented teeth ; lateral spines appearing 
as unpigmented nodes, each surmounted by a transparent peg, but before bleaching 
occurred the lateral spines would, like the teeth, have been strongly pigmented and 
would have been counted as teeth by Newport. Tergites (Fig. 4) : posterior angles 
of T. 9 right-angled, those of T. n and 13 with projections; T. 10 and 12 longer than 
broad with rounded posterior angles. Coxal pores: numerous with no regular 
arrangement. 

Spinulation : 

Ventral Dorsal 

CtPFT CtPFT 

14 a m amp amp a a amp p p 

15 a m amp am a a mp p 

i^th prefemur: slightly expanded distally without any conspicuous setal tuft. 
i$th prefemur: with an ill-defined dorsal longitudinal sulcus and a moderate internal 
distal expansion bearing numerous setae; DpP placed immediately distal to the 
setae. i$th apical claw : simple. Gonopods : short and inconspicuous. 



306 E. H. EASON 

FURTHER SPECIMENS. In addition to the lectotype and three paralectotypes in 
the Museum d'Histoire Naturelle there are five males and two females, dried and 
pinned, in the British Museum (Natural History) labelled "Lithobius elongatus 
Newp." in Newport's hand and "Tunis, 46, 103" on a separate ticket, all of which 
answer essentially to the above description. The same may be said of a female 
from Tunis (B.M.(N.H.) Reg. no. 90.12.16.23) preserved in spirit along with other 
specimens of "L. impressus" collected by Dr. Anderson in North Africa and reported 
on by Pocock (1892). 

None of these specimens has posterior projections on T. 9 and the number of their 
antennal articles varies from 39 to 44. Of variations in spinulation, i4VpT, i4VmT 
and J-5"VpF may be present; i5VaF, i5VaT, i5DpP and i5DpF may be absent. 
Neither i5DaP nor ventral tarsal spines, both of which are found in the next species, 
were present in any of the above specimens. 

17. Lithobius monilicornis Newport 
Fig- 5 

Lithobius monilicornis Newport in Lucas, 1849, p. 384, PL 3, figs 3, 3a, 3b, 30 and 3d 

TYPE LOCALITY. Boudjare"a, near Algiers. 

TYPE SPECIMENS. Syntypes : a male 30 mm long, a rather defective female 35 mm 
long and an immature female 16.5 mm long, all of Eupolybothous impressus (C. L. 
Koch), preserved in spirit, labelled "Lithobius monilicornis Newport, Algerie 
Lucas". Museum National d'Histoire Naturelle. 

REMARKS. Newport distinguished this species from L. elongatus by its more 
numerous antennal articles, smaller prosternal teeth, relatively broader T. 8, 10 
and 12, and longer legs. All these features are shown by the above specimens which 
are undoubtedly identical with the form described from Oran and Algiers by L. 
Koch (1862) under L. impressus C. L. Koch, and we have no reason to doubt their 
identity with L. impressus as originally described from the Algerian coast by C. L. 
Koch (1841) and later figured by the same author (C. L. Koch, 1863). 

More distinctive than the characters given by Newport for distinguishing this 
form from L. elongatus are the posterior projections on T.g (Fig. 5) and the presence 
of ventral tarsal spines on the I4th and i5th legs, both of which were mentioned by 
L. Koch in his description of L. impressus. 

Synonymy of L. monilicornis with L. impressus was first tentatively suggested by 
Meinert (1872). 

DESCRIPTION OF MALE. Size: 30 mm long, 4 mm broad at T. 10, i5th legs 12 mm 
long; if it were not for the contraction of the trunk (Fig. 5) contrasting with the 
extension of that of the lectotype of L. elongatus with exposure of the intersegmental 
membranes (Fig. 4), the difference in length between these two specimens would be 
more marked. Colour: more bleached than the lectotype of elongatus with prac- 
tically no pigmentation. Antennae: 15 mm long with 42 and 43 articles; they 
appear to have been damaged and to have undergone imperfect regeneration; the 
adult female has 49 and 52 antennal articles. Ocelli: not seen owing to bleaching. 



GENUS LITHOBIUS 307 

Prosternum: with 6+6 teeth, faintly pigmented at their apices; being more numerous 
than in elongatus they appear relatively smaller; lateral spines appearing as unpig- 
mented nodes but before bleaching occurred they would, like the teeth, have been 
strongly pigmented and would have been counted as teeth by Newport. Tergites 
(Fig. 5) : posterior angles of T. 9 with small but distinct projections, those of T. 
ii and 13 with more marked projections; T. 10 and 12 broader than long with poster- 
ior angles blunt, not rounded. Coxal pores : numerous with no regular arrangement. 

Spinulation : 

Ventral Dorsal 

CtPFTTa CtPFT 

14 a m amp amp am a a amp p p 

15 a m amp am a a a am p 

14^ prefemur: slightly expanded distally without any conspicuous setal tuft. 
i$th prefemur: with a well-marked dorsal longitudinal sulcus and a prominent 
internal distal expansion bearing numerous setae and a distinct node. i$th apical 
claw : simple. Gonopods : short and inconspicuous. 

FURTHER SPECIMENS. In addition to the three syntypes in the Museum d'Histoire 
Naturelle a female, dried and pinned, in the British Museum (Natural History) 
labelled "L. impressus C. Koch" and "96.3.8.107 Pascoe, Oran" and a number of 
specimens from various localities in Algeria (B.M.(N.H.) Reg. nos. 90.12.16.20-22 
and 24-28; 91.1.18.1-3), preserved in spirit and constituting the majority of the 
specimens of "L. impressus" collected by Dr. Anderson in North Africa and 
reported on by Pocock (1892), all answer essentially to the above description. They 
all have posterior projections on T. 9, the number of their antennal articles varies 
from 43 to 50 and in addition to possessing ventral tarsal spines on the I5th and 
usually also the I4th legs, 15 DaP is always present whereas 15 DpP is found only 
in females. 

DISCUSSION. If the specimens mentioned so far in this paper under L. elongatus 
(see p. 208) and L. monilicornis ( impressus) were the only ones available for examina- 
tion one would have little hesitation in referring the two forms to distinct subspecies 
or even to distinct species, but of Dr. Anderson's North African specimens of "L. 
impressus", those from Constantine (B.M.(N.H.) Reg. no. 91.1.18.4-7) are inter- 
mediate between the two both as regards the shape of the tergites and spinulation, 
so it seems that the characters separating elongatus and impressus are unstable. 
Neither Polybothrus elongatus nor P. elongatus koenigi as briefly described by 
Brolemann (1932) from North Africa can definitely be referred either to elongatus or 
to impressus as defined in the present study, but Lithobius (Polybothrus) elongatus 
var. oraniensis described by Verhoeff (1901) from a number of localities in the 
neighbourhood of Oran seems definitely to belong to L. impressus. L. (Polybothrus) 
impressus corsicus Leger and Duboscq (1903), reported from Corsica and the Mari- 
time Alps (Brolemann, 1930), agrees with impressus in having posterior projections 
on T. 9 and ventral tarsal spines on the i5th legs, but is distinguished by i4DpP 
being replaced by a hook. Of the other six described subspecies, all either from the 
western Mediterranean islands or the European mainland, none can be easily placed. 



308 E. H. EASON 

The species or species group to which all these forms belong is clearly in need of 
revision but, in spite of the intermediate examples from Constantine, it seems 
advisable to retain, for the time being, the distinction between elongatus and 
impressus which belong to the genus Eupolybothrus Verhoeff and the subgenus 
Allopolybothrus Verhoeff as amended by Jeekel (1967) ; but they should be regarded 
as only subspecifically distinct. 

The respective ranges of these two subspecies, if indeed they are true subspecies, 
are difficult to define exactly. C. L. Koch's figure (C. L. Koch, 1963: fig. 105) is 
almost certainly of E. impressus impressus but it was drawn from a specimen from 
Bone which is only 80 Km west of La Calle near the Tunisian border, one of the type 
localities of E. impressus elongatus. The latter, however, seems to be confined in 
North Africa to Tunisia and eastern Algeria whereas E. i. impressus extends along 
the Algerian coast westwards to Oran and probably further into Spanish Morocco. 

18. Lithobius lucasi Newport 

Lithobius Lucasi Newport in Lucas, 1849, p. 385, PI. 3, figs i, la, ib, ic and id 

TYPE LOCALITY. Lac Tonga and Lac Houbeira, Algeria. 

TYPE SPECIMENS. Syntypes: a male 28 mm long, a female 23 mm long and an 
immature male 17.5 mm long, all of L. castaneus, preserved in spirit, labelled 
"Lithobius Lucasii Newport, Algerie Lucas". Museum National d'Histoire 
Naturelle. 

REMARKS. Newport's mention of 5 +5 prosternal teeth, the only character 
appearing to distinguish L. lucasi from L. castaneus, is hardly surprising because, 
although the stout lateral spines which Newport, no doubt, regarded as teeth are 
quite distinct in all three of the above specimens, the 2+2 small teeth are only 
evident in the immature male : in both the adult syntypes the dental margin of the 
prosternum is twisted and distorted giving the impression of a series of projections 
which might be mistaken for teeth. As in the holotype of L. castaneus, the type 
specimens of L. lucasi have oblong coxal pores, lateral spines on the I4th and I5th 
coxae, and the female has a simple claw on the gonopod with only a feeble lobe at the 
base of its external ridge and no denticles: these specimens, therefore, answer to 
Verhoeff's (1942) description of L. castaneus buchnerorum. 

Brolemann (1921) suggested L. lucasi as a possible synonym of L. castaneus: he 
was probably guided by Newport's figure (1849: PI. 3, fig. i) which leaves little 
doubt as to its identity. 

ACKNOWLEDGEMENTS 

My sincere thanks are due to Dr. J. P. Harding and Dr. J. G. Sheals for their 
helpful criticism of my manuscript ; to the Trustees of the British Museum (Natural 
History) for facilities for studying material; to Mr. K. H. Hyatt of the Arachnida 
Section of the Department of Zoology of the Museum for his personal help; to 
Professor G. C. Varley and Mr. E Taylor of the Hope Department of Zoology, 
Oxford, for enabling me to examine specimens from the Newport Collection ; and to 
the Museum National d'Histoire Naturelle, Paris and M. J.-M. Demange for lending 
me the type specimens of Newport's Algerian species. 



GENUS LITHOBIUS 



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E. H. EASON 

BOURTON FAR HILL FARM 

MORETON-IN-MARSH 

GLOS. 



A RE-EXAMINATION OF THE CHESTNUT- 
SHOULDERED WREN COMPLEX OF AUSTRALIA 

By C. J. O. HARRISON 

SYNOPSIS 

THE chestnut-shouldered wren complex within the genus Malurus is re-examined, using recently 
collected material. Seven forms are recognized elegans, pulcherrimus , lamberti, assimilis 
(including master si and bernieri), rogersi, dulcis and amabilis. Variations occur within assimilis, 
rogersi and amabilis. Some variant individuals from the northern edge of the range of assimilis 
show aspects of plumage colour approaching those of other nearby forms. The type of distribu- 
tion appears to provide an example of the refuge concept suggested by Keast. It is suggested 
that pulcherrimus originated in the Eyre peninsula region and assimilis in the Hamersley region, 
and that adaptation to warmer and drier conditions enabled these forms to spread with sub- 
sequent climatic amelioration. A wide tolerance of habitat is shown by assimilis. Other 
forms appear to have more specific preferences but may be occupying the ecological equivalent, 
within their range, of the general habitat required. Evidence of interaction and difference 
between forms indicates that elegans and pulcherrimus behave as good species. 

The taxonomic status of the other five forms appears to be equal, but whether this should be 
specific or subspecific must remain undecided until there is more information on distribution 
and possible interbreeding. 

Certain broad trends in plumage colour and size are apparent. The blue wrens appear to 
have originated as forest birds, probably in the New Guinea region, and to have evolved dull 
female and male eclipse plumages for crypsis. It is suggested that the dull plumage on the 
crowns of breeding males of assimilis may represent a similar trend. 

INTRODUCTION 

When the accumulated specimens of blue wrens, Malurus species, collected during 
the five phases of the Harold Hall Australian Expedition were examined for com- 
pletion of the final report, a re-examination was made of the forms within the chest- 
nut-shouldered wren complex. The last revision had been that of Mack (1934). 
Seven forms are recognized in the present study (Map i), these having been assigned 
various taxonomic ranks during the past. In order to avoid any prior assumption 
of taxonomic status a single specific or subspecific name has been used in referring 
to each population in the following account. In addition to specimens collected 
on the expedition, material already in the collection of the British Museum (Natural 
History) and specimens borrowed from various Australian museums have been used. 

COMMON CHARACTERS 

All birds of both sexes have blue colouring, often rather dull, on the rectrices. 
Males in breeding plumage have a black rump, a broad black collar posterior to the 
ear-coverts and extending round the nape, and black lores. They also have scapular 
patches of rather long chestnut-red feathers, the depth of colour on these tending to 
vary with the intensity of the general plumage colour. The pale edges of the 
tertials are also tinted chestnut-red. When not breeding males moult into an 

Bull. Br. Mus. nat. Hist. (Zool.) 21, 8 



C. J. O. HARRISON 



eclipse plumage similar to that of females and immature birds of both sexes. I have 
used the term "eclipse" plumage in preference to "non-breeding" plumage since 
in some species of Malurus apparently adult males in breeding condition have the 
plain plumage and are capable of breeding while in this plumage; thus while the 
bright male colouring is undoubtedly a breeding plumage the converse is not always 
true. 

The plumage of females, immature birds, and eclipse males is plain brown or 
blue-grey, according to the form. In those forms in which females have chestnut- 
red on the lores and around the eyes, similar colour is present on immature birds of 
both sexes. In these forms adult males do not regain this chestnut-red colour on 
the lores after the first breeding plumage, although there may be some chestnut-red 
on a few feathers immediately around the eye. 

In moulting into the full breeding plumage, males of elegans and pulcherrimus 
appear to acquire the full black lores while still otherwise in eclipse plumage, after 
which bright blue feathers appear first immediately around the eye. The former 
character was not apparent on a large series of specimens of assimilis and a few of 
lamberti and rogersi, save for one brown-plumaged male specimen of assimilis with 
completely black lores. 

Except in amabilis, females and immature birds have reddish-brown bills. Males 
in breeding condition have black bills, but may show some brown on a dark bill 




rogersi 
dulcis 
amabilis 
assimilis 

lamberti 



:::& pulcherrimus 

066688$ 

';> elegans 



MAP i . The ranges as indicated here are very tentative and should be regarded only as a 
generalized diagram for the purpose of discussing relationship. The question marks 
indicate regions where the ranges are in doubt, and within the areas shown populations 
might be small and scattered. 



THE CHESTNUT-SHOULDERED WREN 315 

when in non-breeding plumage; while immature males approaching maturity show 
an increasing spread of black. 

Certain broad tendencies are apparent over the chestnut-shouldered wren group as 
a whole. South-western birds tend to have purple breasts, north-eastern birds to 
have white lores, northern birds to have blue-grey females and eclipse males, and 
eastern birds to be bluer and less violet. Birds tend to become larger towards the 
northerly and southerly limits of distribution. 



FORMS OF THE CHESTNUT-SHOULDERED WRENS 

Seven forms are recognized here and the accompanying table shows the major 
differences in plumage between these. The forms are as follows. 

Lores of ? Plumage of $ Flank of <$ Breast of <$ 

elegans chestnut-red rufous-brown and pale buff blue-black 

greyish-brown 

pulcherrimus chestnut-red olivaceus-brown pale buff dark violet 

lamberti chestnut-red warm brown pale buff black 

assimilis chestnut-red light brown pale buff black 

rogersi chestnut-red light blue-grey white or greyish black 

dulcis white light blue-grey white or greyish black 

amabilis white dark blue very pale buff black 

1. elegans. The Red- winged Wren. 

This is restricted in distribution to the extreme south-west of Western Australia, 
where it is found in thick cover associated with swamps, streams and lakes in areas 
between Gingin and Warriup. 

The male is the palest of these forms, being medium blue on crown and nape, 
gradually changing on the sides of the head to very pale azure blue : paler still on the 
ear-coverts where it has been described as "silvery". The back is an even paler 
and more azure tint. The breast is blackish but with a strong violet tint producing 
a blue-black colour; the latter most apparent when the bird is viewed frontally 
with the bill raised and light falling directly on the underside, the breast appearing 
deep violet with a black band along its lower edge. The hind-flanks and under 
tail coverts are tinted with pale buff. The female is dark greyish-brown on the head, 
and dark rufous-brown on the back and wings. Below it is light greyish-buff on 
throat and breast and pale buff on belly and flanks. The lores are deep chestnut-red. 

2. pulcherrimus. The Blue-breasted Wren. 

This form appears in a zone north of that of elegans and mostly south of that of 
assimilis. Its western limits are between the mouths of the Murchison and Namban 
Rivers and its range extends through the mallee and wheatbelt in a south-easterly 
zone to Warriup and Eucle, with an apparently isolate population on the Eyre 
Peninsula. 

The male is a deep violaceous blue on the head and deep violet on the back. The 
forehead and sides of the head are more blue, becoming light blue on the ear-coverts. 



316 C. J. O. HARRISON 

The breast is glossy dark violet, brighter towards the edges of the sides, and with a 
narrow black band along its lower edge. The flanks and under tail coverts are a 
drab light buff. The female is dull olivaceous brown, with chestnut-red lores. 
Below, the throat and breast are pale greyish-buff, the belly white, and the flanks 
and under tail coverts light buff. 

3. lamberti. Usually regarded as the eastern form of the Variegated Wren, 
assimilis; but sometimes called Lambert's Wren. 

This form occurs on the eastern seaboard, and the range appears to be the region 
east of the Great Dividing Range, south to Sydney and north at least to the Brisbane 
region, although there appears to be little evidence of what occurs near the coast 
north of this. Further inland, specimens collected by Elsey (Macdonald and Colston 
1965) on the Belyando River are certainly assimilis and the latter is said to have 
been collected on the Dawson River (White 1916). These rivers, although inland 
in the complex topography of the Dividing Range in mid-Queensland, drain towards 
the east. Two female specimens collected at Bloomsbury, near Prosperine, on the 
Wilkins Expedition (allowing for foxing and comparing them with material taken 
elsewhere on the same expedition) are of the lamberti form, suggesting that the 
latter extends well up the east coast. 

Males of this form are deep blue, but not violet-blue, on the back and nape; 
becoming paler blue on forehead, sides of head and ear-coverts, the last having, in 
comparison with other light blue plumage, a slight azure tint. There are violet 
tips to feathers bordering the sides of the black breast. Posterior flank feathers are 
pale buff. Females, immature birds and males in eclipse are brown above, tinted 
with warm buff on the rump; pale below with yellowish buff on flanks and belly; 
and have deep chestnut-red on the lores and a narrow ring round the eye. The 
brown colour on these birds is darker and warmer in tint than that of assimilis. 

4. assimilis. The, Variegated Wren. 

This form appears to occur from the Great Dividing Range and its ancillary ranges 
in Queensland, westwards across the entire dry central region to the west coast, 
north to the Gull of Carpentaria, the Roper River, and the Broome Area of Western 
Australia, and south to the edges of the range of pulcherrimus but apparently not 
as far as the south coast. 

Males are violet-blue on the back, nape and crown, grading into deep blue on the 
forehead and light blue, often slightly azure, on ear-coverts and around the eye. 
Some individuals are less violet dorsally, lighter blue on the head, and more azure 
on the ear-coverts. The posterior flanks are pale buff. There are violet tips to 
feathers bordering the sides of the black breast. Of 33 adult males apparently in 
full breeding plumage and not moulting, 17 showed an area of dull brownish colour 
on the crown on the head, often resembling a distinct but irregular cap. Another 
five show traces of this and only n have fully-coloured heads. Females, immature 
birds and males in eclipse are a dull, light brown above, with a buffish rump. 
Below they are pale buff, deepest on the flanks and almost absent from the throat 
Adult males in this plumage, in addition to lacking the chestnut-red lores present on 
the others, are much whiter below. 



THE CHESTNUT-SHOULDERED WREN 317 

5. rogersi. Usually regarded as the Western form of the Dulcet or Lavender- 
flanked Wren, dulcis. 

From specimens collected, and from others kindly lent by the Western Australian 
Museum, this form occurs in the ranges of the Kimberleys from the Leopold Range 
north to Napier Broome Bay and westward to the Ord River. 

Dorsally males of this form are virtually indistinguishable from those of assimilis, 
but do not show the extreme violaceous tint of some individuals of the latter and 
tend to come about the middle of the range of variation of the blue and violet colours. 
There is some individual variation. The wing feathers, both flights and coverts, 
are a darker brown than those of assimilis and may show a faint bluish sheen. 
This is apparent in fresh plumage, but an otherwise moulted male specimen showed 
old wing feathers of a similar, sandier brown colour to those of assimilis, with just 
a few darker new feathers. There are violet tips to feathers bordering the breast. 
The belly and flanks are white, and of ten specimens only one shows a faint greyish 
wash with even fainter violet tint which might have given rise to the vernacular 
name. In the circumstances it would seem more correct to call this species "White- 
flanked "rather than "Lavender-flanked" if it is desirable to refer to this aspect of the 
plumage. 

Females, immatures and males in eclipse plumage all have a dorsal plumage of 
dull blue-grey, becoming paler blue on the sides of the head and neck. Males are 
more blue and less grey, and very young birds are greyer, with a hint of brown, and 
less blue. Wing-coverts are dark brown with greyish edges. Both females and 
immature males show the chestnut-red lores that substantiate Mathew's rogersi. 
Females show a very pale buff tint on the underside, while males in eclipse plumage 
are almost white below. 

6. dulcis. The typical Dulcet or White-flanked Wren. 

This form is usually said to have a rather restricted distribution between the 
Mary and King Rivers (the northern King River), in Northern Territory; (Storr 
1967). Humphries (1947) refers to M. amabilis [=dulcis~\ being observed on one 
occasion at Melville Bay, and more recently Rix (1970) has recorded this species at 
Elsey Creek on the upper Roper River. It therefore seems possible that its true 
range may be around, or through, the Arnhem Land Region. 

Males appear almost indistinguishable from rogersi but in fresh plumage appear to 
show a more obvious bluish wash on the wing feathers, particularly the coverts, 
making these appear darker in colour. Females and immature males differ from 
rogersi in being slightly darker dorsally, but more conspicuously in having the lores 
and a narrow ring round the eye creamy-white and not chestnut-red. Ventrally 
the very pale buff colour is apparent but is combined with a faint grey tint to give 
the plumage of the underside a rather drab appearance. 

7. amabilis. The Lovely Wren. 

This is another form with apparently limited distribution, occupying habitats 
bordering the rainforest along the north-east Queensland coast between Cape York 
and Cardwell, but it has also been recorded (Thompson 1935) from the Gulf coastal 



318 C. J. O. HARRISON 

regions on the Lower Edward River where it was noted and collected in "dense dry 
scrubby country on raised beaches". 

Males are most similar to lamberti, but are a lighter, clearer blue on the back and 
head, without any definite violet or azure tints, the ear-coverts being similar in 
colour to the rest of the head. The violet tips to feathers at the sides of the breast are 
restricted to a few vestigial tips. The flanks show some pale buff. The pale edges 
to wing feathers show a more distinct blue wash, and these feathers are otherwise 
very dark in colour, in some instances practically black, and in the case of the 
lesser coverts usually black. 

Females, immature males and males in eclipse resemble those of dulcis in having 
creamy-white lores and eye-rims, but differ in the much darker colour of the dorsal 
plumage. The latter is a dark and rather dull blue, becoming a little lighter on the 
forehead, and with conspicuous light blue ear-coverts. Both females and males in 
eclipse show black bases to the feathers across the upper mantle which are wholly 
black in the breeding males. The bill is black in both sexes. Wing feathers are 
dark with a dark blue wash. Below the plumage is very pale buff, whiter on the 
throat, and whiter overall on males. The juvenile is browner and less blue on the 
back, and has a dark, blackish-brown bill. 

VARIATION WITHIN FORMS 

Individual variation is apparent within assimilis, rogersi and amabilis. It is 
assimilis, with its very extensive range, which shows the greatest variation. Indi- 
viduals showing some variation were described as new races and originally nine were 
recognized within the range of this form, but these were reduced by Mack (1934) to 
three (Map 2). On the basis of the material examined, I cannot agree that there 
are three readily recognizable subspecies within this area, but there is some tendency 
toward broad trends in colour variation within the very large range occupied by 
this form. 

As already mentioned, the only obvious difference is for some individuals to have 
paler heads. In comparing the blue colour it is necessary to have specimens side 
by side at the same angle to the light, since the incidence of light can affect the 
apparent hue of structural colours of feathers. 

The original description of assimilis was based on one of the darker south-eastern 
specimens. Grant (1909) described bernieri, from Bernier Island off the west coast, 
differing in that the ear coverts of the male were a dark blue. In a specimen examined 
they are undoubtedly dark, but match well with those of a specimen from near 
Lake Frome in South Australia, and one from near Hughenden in northern 
Queensland. The third race, mastersi was described by Mathews (1912) from a 
specimen from Alexandra, Northern Territory, with more azure blue ear-coverts. 

If a dark specimen is compared with a light one the difference is quite obvious, 
but it was found possible to lay out a series of adult males which showed a consistent 
gradation of forehead and ear-covert colour, from one extreme to the other, with 
no division at any point to suggest that more than one population was involved and 
there was no clear geographical pattern. The specimens, beginning with the deepest 



THE CHESTNUT-SHOULDERED WREN 319 

colour, were from the following localities: i, South Australia (unlocalized older 
specimen, probably from south-east settlement); 2, Bernier Island, W.A. ; 3, 30 
miles west of Lake Frome, S.A. ; 4, near Hughenden, Qld; 5, 15 miles west of Lake 
Frome, S.A. ; 6, Bourke, N.S.W ; 7, Finke River, N.T. ; 8, Tambrey, W.A. ; 9, Point 
Cloates, W.A. ; 10, Prairie, Qld; n, Little Desert, Vic,; 12. Warburton Mission, 
W.A. ; 13, Tambrey, W.A.; 14, Upper Gascoyne, W.A. ; 15, Moonlight Creek, Qld.; 
16, Upper Gascoyne, W.A.; 17, Fitzroy River, W.A. ; 18, Upper Gascoyne, W.A. 
(see map 2). 

If one were to follow Mack (1934) nos. 6 and n would form his dark subspecies; 
no. 2 another; nos. i, 3, 5, 7-9. 12-18 would form a pale subspecies; and nos. 4 and 
10 would be part of an indeterminate population. Mack himself appeared uncertain 
of the precise limits of the forms which he recognized and in his map of subspecific 
distribution (map 2) did not show precise areas but indicated lobes from a main 
mass. Condon (1951), in discussing South Australian birds, abandoned head and 
back colour as diagnostic characters and disagreed with Mack's views on subspecific 
limits. Ford (1966) noted the variability of the head colouring in Western Australian 
birds. 

There is some evidence of a general tendency for birds with deeper and darker 
blue colour to occur on the eastern and extreme western edges of the overall distri- 
bution, and for paler birds to be most frequent in the desert areas of Western Aus- 
tralia and towards the north-western limits of distribution. Perhaps Gloger's rule 




MAP 2. The dotted line indicates Mack's version of the distribution of races of M. lamberti : 
A, M. I. lamberti; B, M. I. assimilis; c, M. I. master si; D, M. I. bernieri. The figures are 
those indicating individuals in the section on variation within forms. The black dots 
indicate the localities of variant individuals. 



320 C. J. O. HARRISON 

is involved. This clinical trend does not, however, appear to allow convenient 
subdivision and, in addition, there is evidence of variability between individuals in 
small areas as well as over a wide range. Nos. 8 and 13 from the above list were 
collected on the same occasion from the same party of birds, and there are similar 
variations within specimens from single localities at Fitzroy River, Western Australia, 
and in northern Queensland. T. Carter (Mathews 1922-3) noted a bird taken on the 
Minilya River as "blue" on the back, although most individuals from the west 
coast area were distinctly violet. 

Other characters for separation have been suggested. Condon (1951) stated that 
master si could be distinguished by its clear white abdomen. This is difficult to 
determine in the skins of very small birds prepared by a variety of hands, but it 
was not apparent in the material now examined (46 males) and where some slight 
difference was apparent it did not relate to the earlier subspecific divisions. 

Condon and others also refer to differences in the relative lightness of the brown 
colour of the wings. The glossy black and blue body plumage appears to be stable 
but the brown plumage of the wings shows a definite tendency to fade on the living 
bird and most of the variation apparent in specimens examined due to the differing 
age of the feathers, the fresh new feathers being darkest. In addition there is some 
change due to foxing in older specimens. The wing moult and body moult do not 
appear to be closely synchronised, and birds which show complete breeding plumage 
on the body may still be actively moulting and growing wing feathers. The relative 
depth of colour on the wings does not therefore provide a consistent and useful 
taxonomic character. 

The difference in plumage between individuals within assimilis are so slight and 
gradual that there seems no reason to suppose that more than one unit is involved 
with some local differentiation beginning to occur. I am of the opinion that this 
should be treated as a single form, and assimilis North, 1901, is the oldest name; 
mastersi Mathews, 1919, and'bernieri Grant, 1909, being synonyms. 

MAJOR VARIATION IN THE RED-SHOULDERED WREN GROUP 

Among specimens of rogersi a male from Kulumburu (W.A. Museum, A 8884) 
was a much lighter blue on the head, and similar in this respect to a specimen of 
assimilis (B.M. no. 1964. 60.585) from Moonlight Creek, north Queensland. 

There is some variation in the blue tint of the head of breeding males of amabilis. 
Mack (1934) separated birds from the southern half of the range as a subspecies, 
clams, on the grounds that they showed lighter blue colouring. An examination 
of specimens, including some lent by the Queensland Museum, revealed a small 
difference between extremes. Birds recently collected from Tully were a lighter 
blue than some early specimens from Somerset and Port Albany, and the latter showed 
a slight violaceous tint to the deeper blue which was a little more distinct on some 
northern birds from the Queensland Museum. Recent specimens from Ayton, 
towards the middle of the range of this form, appeared to be intermediate in character. 
The total difference appears to be small and clinal in character and on the material 
at present available I would regard clams Mack, as a synonym of amabilis. 



THE CHESTNUT-SHOULDERED WREN 321 

VARIANT INDIVIDUALS 

In addition to the variations already described there are some variant specimens 
which, from their appearance and locality, (map 2) may throw some light on the 
relationship between various forms. 

The most conspicuous of these is an adult female assimilis (B.M. no 1964.60.578) 
collected on the Norman River south of Normanton, Queensland, at the south- 
east corner of the Gulf of Carpentaria. This bird shows a pale greyish tint over the 
whole body plumage, particularly noticeable on the sides of the head and neck. 
The lores are the normal chestnut-red colours and the retrices dull blue. There is 
no suggestion that this variation is due to wear or fading. Normal brown plumage 
contains both eumelanin and phaeomalenin and these are not differentially affected 
by exposure to light, (Harrison 1963). The bird was collected from a party of 
eight individuals and another female and an immature male collected at the same 
time appear to be normal. 

Two other females of assimilis (B.M. nos 1969.4.415, 417) show similar, although 
less marked, tendencies towards greyish plumage. These are the only two females 
taken on the Roper River, Northern Territory (14 I5'S., 135 3' E.). On these 
the grey tint is superimposed on the normal brown plumage to give a colder, greyer 
tint, but the sides of the head and neck are noticeably grey, and the rump has an 
olive tint. 

A minor plumage variation found during close examination was the presence of a 
few dark lesser covert feathers on some males of assimilis, the coverts of this form 
normally being brown. Occasional blackish covert feathers occurred on a male 
(B.M. no. 1964.60.674) from Norman River, at the same "locality as the female 
described above, but from a different party of birds; and on a male (B.M. no 1964. 
60.585) from the south of Moonlight Creek, on the south-east of the Gulf of Carpen- 
taria. Males showing some blue tips to feathers occurred at Moonlight Creek 
(B.M. no 1964.60.581), Roper River (1969.4.414, 416), and at Mount Anderson 
(1964.4.443, 450) in the south-western Kimberleys. 

These variants may be significant in view of their distribution. The greyish 
females of assimilis occur on the edge of its range, in one case near amaUlis and in 
the other near dulcis, both of which have blue-grey females. The variant wing- 
coverts similarly occur on individuals on the northern edge of the range, birds with 
blackish coverts at Norman River and Moonlight Creek being near amabilis which has 
similar coverts ; and birds with blue covert tips at Roper River and Mount Anderson 
being near to dulcis and rogersi respectively, both of these having blue on the converts. 
The exception is the second male from Moonlight Creek which has the few covert 
tips blue and not blackish. 

ZOOGEOGRAPHICAL DISTRIBUTION PATTERN 

The type of distribution shown by the chestnut-shouldered wren complex appears 
to provide a good example of the refuge concept suggested by Keast (1961) and to 
show the utilisation of a greater number of refuges than do the other superspecies 
groups which he instances. Keast did use these birds as an example but used the 
races as mapped (map 2) by Mack (1934) and hence found the parallels with other 



322 C. J. O. HARRISON 

species groups less obvious than they should have been. There is, in fact, a ring of 
forms, most of which still occupy limited peripheral areas (map i). On the eastern 
edge of Australia is lamberti, amabilis is in the north-eastern peninsula, dulcis in the 
north of Northern Territory, rogersi in the Kimberleys of the north-west, and 
elegans in the south-west corner. Since pulcherrimus is present on the Eyre Penin- 
sula as well as in the south-west it is more likely that, rather than having evolved 
in competition with elegans, it differentiated in or somewhere near the former region 
in a refuge demanding tolerance of drier conditions and higher temperatures than 
did the south-west refuge; and therefore, when conditions ameliorated, it would 
have been able to spread westwards into similar habitats bordering the range of 
elegans. 

The last form, assimilis, presents a slight problem. There are several potential 
refuges which might have been available (Keast, 1961) and are not occupied by 
other members of this group, but from its present distribution it is reasonable to 
assume that the differentiation would have occurred in a refuge where conditions 
were both drier and warmer than in some others. The extreme south-east is there- 
fore unlikely since conditions there are likely to have been both moister and cooler. 
The two remaining likely areas are the Hamersley region of coastal Western Australia 
and the central ranges. The differentiation in isolation apparent in other taxa 
(Keast 1961) occurs less frequently, in the latter area and the Hamersley region 
would seem the more likely of the two. The adaptation to such a refuge would 
give this form the slight advantage that would enable it to exploit more rapidly and 
successfully the gradual amelioration of extremely arid conditions of the central 
area and to spread to produce the apparent "Eyrean" distribution (Spencer 1898). 
Keast (1958) has pointed out that many forms with this distribution are derived 
from western isolates of species with more extensive distribution at an earlier period. 

HABITAT PREFERENCE 

From the limited distribution of some of these forms, they may have evolved 
some degree of habitat preference which might limit any subsequent spread. In the 
south-west elegans has a limited distribution in low cover bordering fresh water 
swamps or streams; while pulcherrimus is a species of sandplain scrub and mallee. 
Of the northern forms, both rogersi and dulcis occur in regions of ranges and plateaus 
and appear to be mainly confined to places where the floors and sides of sandstone 
or granite gorges have low scrubby vegetation and where natural breaks or dis- 
continuities in the rock of the ranges are accompanied by vegetation including low 
bushes and spinifex. The other extreme is shown by amabilis which occurs on the 
outer edge of rain-forest or in suitable low cover in open forest adjoining it. The 
information on lamberti is poor but it appears to occupy thick, shrubby growth in 
fairly moist habitats including the thick shrub layer of forest, and in this respect 
would seem to show some similarity to amabilis. 

As might be expected from its considerable range, assimilis shows a wider habitat 
tolerance. In general it tends to be a bird of shrubby growth bordering water- 
courses in drier regions. It may, however, move into the sparse vegetation of 
sandstone ridges, occupying a similar habitat to rogersi and dulcis in similar regions, 



THE CHESTNUT-SHOULDERED WREN 323 

and although from their field observations B. M. Booth and D. Freeman are of the 
opinion that in such places it is less likely to venture onto bare rock than are the last 
two forms this would not prevent it from sharing the same cover were it to occur 
with them. At the other extreme it extends into the shrub growth along rivers and 
creeks in open forest; and down into thickets, riverine forest and the edges of 
mangroves. 

In Western Australia near Carnarvon, where this form occurs in the scrub on 
dunes among saltbush, pairs were seen out in the mangroves apparently disputing 
territory over rising seawater. The mangroves in this area form a narrow belt 
separated from the beach by a tidal lagoon up to about half a mile wide in places. 
Within the mangroves, which are low and open on the landward side, becoming 
taller and thicker to seawards, there are small crescentic beaches with a sparse 
growth of herbaceous plants. Although it seems likely that the presence of pairs 
in such a place may have been due to a lack of suitable territories due to overpopu- 
lation, and the fact that these were pairs rather than parties suggests that they were 
young or breakaway units searching for new areas, their presence nevertheless indi- 
cates the readiness of this form to attempt to occupy a wide range of habitats. 

This wide habitat tolerance in assimilis does suggest that the ecological require- 
ments of the birds may be relatively simple and that, at least where the northern forms 
are concerned, the apparently narrower habitat preference of various other forms may 
be simply due to the fact that they are occupying the ecological equivalent within a 
more specialised and less varied biotope. Certainly the habitat tolerance of assimilis 
is such that were it to come into contact with other forms it seems likely that it 
would occupy the same niche, and the apparent minor variation in habitat would be 
unlikely to act as a barrier between them. 

THE INTERACTIONS AND TAXONOMIC STATUS OF THE FORMS 

Where a number of similar allopatric forms exist it is always difficult to determine 
their precise taxonomic relationship. In the present instance the recognition of 
five species for the seven forms appears to be an accidental result of the successive 
description of the forms and their similarity to those already known at the time of 
their discovery. 

The situation in the south-west has been well investigated (Serventy 1951, Ford 
1966, 1969). Here elegans and pulcherrimus have contiguous ranges and pulcheri- 
rimus and assimilis are sympatric in the north-west part of the former's range. 
Ford (1966) has evidence that these pairs encounter each other in the field; but 
elegans and assimilis do not meet. In such encounters these three forms appear to 
ignore each other and behave as good species. Where pulcherrimus and assimilis 
are sympatric they occur in the same biotope and show a mosaic distribution, but 
it is not certain whether this indicates interspecific intolerance or differing responses 
to microhabitats (Ford 1966). 

The plumage of breeding males appears adequate to ensure specific recognition, 
the combination of colour on breast, head and back being conspicuously different in 
the three (table i). A combination of pale azure and blue-black is present on 
elegans; violet-blue and dark violet on pulcherrimus; and lighter violet-blue and 



324 C. J. O. HARRISON 

black on assimilis. These colours would be particularly conspicuous in any frontal 
displays. The interspecific variation in colour of females, young and eclipse males is 
relatively slight, but this might aid specific recognition. There may be other 
differences. Ford (1966) refers to a detectable difference in the voices of pulcherrimus 
and assimilis. Size differences are relatively slight, with a clinal increase in a 
south-westerly direction; and I doubt if the differences in bill-size given by Ford 
would be sufficient to produce the variation in food selection which he suggests. 

The situation is more complicated in the northern half of Australia where there 
are five recognizable forms lamberti, assimilis, rogersi, dulcis and amabilis pre- 
viously recognized as three species lamberti I assimilis, roger si j dulcis and amabilis. 
These five forms show slight size variation of a clinal type, the more northerly 
being a little larger. Since within the whole chestnut-shouldered wren group 
size tends to increase towards the northern edge of the distributional range as well 
as the southern edge it seems inadvisable to suggest the latter as an example of 
Bergmann's rule. 

Apart from this, the characters which differ between forms and could be used for 
recognition are those of plumage colour. In Malurus species generally specific and 
subspecific variations are usually most apparent in the breeding plumage of the 
male. In the present group although the dorsal blue colour varies, the range of 
variation within assimilis encompasses that of both rogersi and dulcis. In sequence 
from blue to violet-blue the arrangement would be amabilis lamberti assimilis 
(inc. rogersi and dulcis) ; the difference between the last two of the three, usually 
regarded as conspecific, being greater than that between assimilis and the forms in 
parentheses which are regarded as good species. 

The only other obviously variable plumage character of breeding males, which 
has been used for separation of forms, is the flank colour (table i). The posterior 
flanks show a variable amount of pale buff on lamberti and assimilis, slightly buff 
colouring on amabilis, and are usually white, with a purplish-grey wash on some 
individuals of rogersi and dulcis. One or two specimens of the last form show a 
small amount of pale buff, particularly around the upper thighs, but since buff is 
present on females and eclipse or immature males, this may only indicate incomplete 
assumption of full male plumage. 

The female plumage shows greater distinctiveness in this group, varying between 
brown and blue-grey ; lamberti being warm brown, assimilis normally a paler, duller 
brown, rogersi and dulcis pale blue-grey, and amablis deep blue-grey. The apparent 
clear-cut distinction is blurred a little by the existence of the greyer variant individ- 
uals of assimilis. If these are included in the sequence there is a much smoother 
gradation and the most relevant differences would appear to be the darker colour of 
amabilis and the change from chestnut-red lores on lamberti, assimilis and rogersi to 
white lores on dulcis and amabilis. 

The mainly allopatric distribution makes it difficult to judge the amount of 
interaction that might potentially occur between these northerly forms. Mack (1934) 
suggested that assimilis and lamberti interbred in northern Queensland, but from 
his account it seems possible that he did not make allowance for variation within 
assimilis, and a more satisfactory investigation of this is still needed. At the oppos- 



THE CHESTNUT-SHOULDERED WREN 325 

ite extreme of the range of the latter the fifth phase of the Harold Hall Australian 
Expedition found both assimilis and rogersi near Mount Bell in the Leopold Range, 
one of the more level plain and the other in the gorges of the range, but in types of 
habitat in which they might well encounter each other. Intergrades between the 
two would be very difficult to identify. Assuming that interbreeding produced 
individuals with intermediate plumage colour, this would only involve the buff 
on the flank of the male and the body colouring of birds other than breeding males. 
Any male bird with buff on the flanks would probably pass as assimilis. Inter- 
mediates when female or immature should be recognizable and would probably 
resemble the greyer variant individuals of assimilis already described. If these 
single plumage characters were controlled by a simple pair of dominant and recessive 
alleles recognition of a hybrid might be impossible. 

Rix (1970) has recently recorded a party of assimilis in a clump of teatree on 
Elsey Creek with a party of dulcis (referred to by Rix as amabilis) in an area of tall 
grass with a few small shrubs only about twenty yards away. There is therefore 
a potential zone of interaction between assimilis and dulcis also; but theoretically 
none between assimilis and amabilis (map i.) However, the presence of variant 
individuals of assimilis apparently showing the plumage character of the other 
two to some degree, at points in its range nearest to the known ranges of the 
other two forms, suggests either that the factors responsible for these plumages 
are environmental ones which may act upon individuals of assimilis in the same 
regions and tend to select for similar characters (although in such circumstances one 
might expect all individuals in an area to show some evidence of this) or that there 
is some interbreeding within the areas where the forms approach each other. If the 
latter is true then it would suggest that the distribution of amabilis is more extensive 
than our present knowledge indicates. To suggest such interaction presupposes that 
there are not fully effective barriers of ecology or species-specific recognition between 
the forms. The obvious ecological barrier of habitat preference has already been 
discussed and it is suggested that the wide habitat tolerance of assimilis could 
potentially bring it into contact with the other forms. Recognition would be based 
on signals of behaviour, voice or plumage pattern. Such little evidence as we have 
suggests that these various forms are similar in their behaviour and that their 
voices are indistinguishable, although in view of Ford's (1966) comment on the voices 
of two south-western forms a more critical appraisal of this character might be 
helpful. 

In the south-western forms, where plumage colour of the breeding males appears 
to be a good specific character, adjacent forms show marked variation in the com- 
bined colouring of head, back and breast. This suggests that such signal colouring 
is associated with frontal displays, and such postures have been described (Rowley 
1964) in the Superb Blue Wren, M. cyaneus, the only adequately studied species. 
The present forms all have black breasts, with slight variation in the extent of the 
violet edge at either side. The head and back colouring is indistinguishable and 
variable in three forms and it is therefore difficult to argue that the slight differences 
in the shade of blue shown by the other two is of any great significance. The 
flank colour used to separate forms would not appear to be used in displays and, 



326 C. J. O. HARRISON 

in some at least, might be actually concealed by the wings. There would therefore 
seem to be no reason to regard the male breeding plumage as obviously specifically 
isolating. 

An alternative isolating factor would be the recognition by the male of differences 
in the female colour, the grey or brown plumage, the presence or absence of chestnut- 
red lores, or the bill colour. Heterogynism variation in females but not in males 
occurs in races of some species but is comparatively rare. Mayr (1942) records it in 
three races of Pachycephala pectoralis in the Solomons, but since these are on different 
islands the question of recognition or interaction would presumably not arise. 
Hellmayr (1929) and Zimmer (1931 et seq.) also found it in races of some species of 
Formicariidae in South America, but the implication of this in relation to recognition 
does not appear to have been examined. There is therefore no useful information 
on the use of heterogynism in intraspecific recognition. Encounters of forms in 
which chestnut-red lores are present with those which lack them might produce some 
confusion since in the forms that possess them the absence of these usually indicates 
that a bird in brown or blue-grey plumage is an adult male in eclipse. Similarly 
a black bill on a bird in eclipse plumage usually indicates a male, but females of 
amabilis also have a black bill. 

In summary, there are two very distinct forms of the chestnut-shouldered wrens 
in the south-west which can be regarded as separate species; but the inter-relation- 
ships of the remaining five forms are obscure. Each has a slightly different com- 
bination of plumage characters. The selection of particular characters for the pre- 
vious separation of species seems to have been arbitrary, and each form would appear 
to represent a separate isolated evolutionary unit. The more recent taxonomic 
treatment of the peripheral rings of isolate forms, of which there are many in the 
different taxa on the Australian sub-continent, has been to regard these as species, 
except where secondary re-integration occurs in forms which spread and re-encounter 
each other, in which case they are regarded as subspecies. A typical example of 
the latter is Keast's (1961) interpretation of relationship in the Australian forms 
of Sittella. 

In the case of the chestnut-shouldered wrens there are five distinct forms of 
equal standing, all of which could be regarded as specific or near-specific entities. 
It is possible however that speciation has been incomplete and that they might 
interbreed and re-integrate freely if they came together again. The extensive 
spread of assimilis has increased the likelihood that this might occur, and the 
existence of variant individuals suggests that it may be taking place. If subsequent 
field investigation shows that this is so, and if more detailed study of these forms 
confirms that there are no barriers, of the type discussed, to prevent this occurring, 
then it might be preferrable to regard them as subspecies within a single species. 

APPARENT EVOLUTIONARY TRENDS 

The genus Malurus appears to have affinities with a group of genera in New 
Guinea (Harrison and Parker 1965, Harrison J_969a). The deep blues, black, and 
chestnut-red of these birds is a colour combination which appears more usually to 
originate in moist tropical forest; and from this, and from what we know of the 



THE CHESTNUT-SHOULDERED WREN 327 

general zoogeographical trends within this region, it seems probable that Malurus 
as a unit may have invaded Australia from this direction and subsequently evolved 
and radiated within the Australian environment. In the New Guinea group as a 
whole females are similar in plumage colour to males, or as richly coloured. The 
production of the rather drab female plumage, and of an eclipse plumage in the non- 
breeding males appears to have occurred in Australia, and it would seem most 
likely that this is an adaptation for crypsis in a generally more open environment 
where the more brightly coloured bird is vulnerable to predation. In the case of 
the Blue-and- White Wren, M. leucopterus, there appears to have been a suppression 
of bright male plumage in subordinate males (Harrison 1969^, the alternative 
possibility being that of very high predation on males in breeding plumage. The 
species is one that occurs in arid areas with relatively sparse cover, and the blue and 
white male is very conspicuous. 

In view of this apparent tendency to reduce the conspicuous colour of the males of 
this taxon in the more open areas, retention of the dull eclipse plumage on the crowns 
of so many of the collected male specimens of assimilis may be significant. With 
such a bird in its normal posture with tail cocked acutely and head back a little, 
the back is relatively less conspicuous and to an aerial avian predator the crown 
of the head must be the most obvious mark. A dull crown might thus carry a strong 
selective advantage. 



ACKNOWLEDGEMENTS 

I am very grateful for the loan of material to the Western Australian Museum, 
the Arid Zone Research Institute of Northern Territory, the C.S.I.R.O. Division of 
Wildlife Research at Canberra, and the Queensland Museum. I should also like 
to acknowledge the help and information that I have received from Major B. M. 
Booth, Mr. D. J. Freeman, Mr. C. Frith, Mr. I. C. J. Galbraith, Mr. S. A. Parker, 
Dr. D. W. Snow and Dr. G. M. Storr. 



REFERENCES 

CONDON, H. T. 1951. Notes on the birds of South Australia; Occurrence, distribution and 

taxonomy. S. Aust. Orn. 20 : 26-28. 
FORD, J. 1966. Taxonomy and variation of the Chestnut-shouldered wrens of Western 

Australia. Emu 66 : 47-57. 

1969. Range limits of the Blue-breasted Wren in Western Australia. Emu 68 : 283-284. 
HARRISON, C. J. O. 1963. Grey and fawn variant plumages. Bird Study 10 : 219-233. 
ig69a. The affinities of the Blue Wren genus Malurus and related genera; with special 

reference to the grass-wren genus Amytornis. Emu 69 : 1-8. 

i96gb. Helpers at the nest in Australian passerine birds. Emu 69 : 30-40. 
HARRISON, C. J. O. & PARKER, S. A. 1965. The behavioural affinities of the Blue Wrens of 

the genus Malurus. Emu 65 : 103-113. 
HELLMAYR, C. E. 1929. On heterogynism in Formicarian birds. /. Orn. Lpz., Ergbd. 

2 : 41-70. 
HUMPHRIES, C. P. 1947. Among the birds at Melville Bay. Emu 47 : 130-136. 



328 C. J. O. HARRISON 

KEAST, A. 1958. The genus Psophodes Vigors and Horsfield, and its significance in demon- 
strating a possible pathway for the origin of Eyrean species from Bassian ones. Emu 
58 : 247-255. 

1961. Bird speciation on the Australian continent. Bull. Mus. Comp. Zool. Harvard 

123 : 303-495- 

MACDONALD, J. D. & COLSTON, P. R. 1965. J. R. Elsey and his bird observations on Gregory's 
overland expedition, Australia, 1856. Emu 65 : 255-278. 

MACK, G. 1934. A revision of the genus Malurus. Mem. Nat. Mus. Viet. 8 : 100-125. 

MATHEWS, G. M. 1922-23. Birds of Australia. Vol. 10. Witherby; London. 

MAYR, E. 1942. Systematics and the origin of species. Columbia U.P. : New York. 

OGILVIE-GRANT, W. R. 1909. [Three new species of birds from West Australia]. Bull. Brit. 
Orn. Cl. 23 : 72. 

Rix, C. E. 1970. Birds of the Northern Territory. South Australian Ornithologist 25 : 147- 
190. 

ROWLEY, I. 1964. The life history of the Superb Blue Wren, Malurus cyaneus. Emu 
64 : 251-297. 

SERVENTY, D. L. 1951. The evolution of the Chestnut-shouldered Wrens (Malurus}. Emu 
51 : 113-120. 

SPENCER, B. 1898. Report of the work of the Horn scientific expedition to Central Australia. 
Narrative. Dulau: London. 

STORR, G. M. 1967. List of Northern Territory Birds. Spec. Publ. W. Austr. Mus. No. 4. 

WHITE, H. L. 1916. A new Wren-Warbler, Malurus lamberti dawsonianus (Barnard's Wren). 
Emu 16 : 69. 

ZIMMER, J. T. 1931 et seq. Notes on Peruvian birds. Amer. Mus. Novit. 500 et seq. 



Dr. C. J. O. HARRISON 

Department of Zoology 

BRITISH MUSEUM (NATURAL HISTORY) 

CROMWELL ROAD 

LONDON, S.W.7 



A NEW SPECIES OF LIRONECA (ISOPODA; 

CYMOTHOIDAE) PARASITIC ON CICHLID 

FISHES IN LAKE TANGANYIKA 

By R. J. LINCOLN 



INTRODUCTION 

ONLY three species of parasites belonging to the family Cymothoidae have been 
recorded from freshwater localities on the African continent. The first of these to 
enter the literature was Ichthyoxenus expansus, described by Van Name in 1920 as a 
parasite on the gills of the citharinid fish Eugnathichthys eetweldii Boulenger. This 
isopod has since been widely recorded from the waters of the Congo river basin 
(Monod, 1931 ; Dartevelle, 1939; Brian & Dartevelle, 1949 and Grosse, 1963). There 
has been much discussion in the past about the validity of the genus Ichthyoxenus 
to which expansus was referred. Miers (1880) expressed the opinion that the two 
genera Ichthyoxenus and Lironeca were indistinguishable, except that the former 
was exclusively freshwater. Van Name (1920) considered the wide separation of 
the localities from which the species of Ichthyoxenus had been described as evidence 
that they were not a monophyletic group, but were instead convergent forms. In 
this view the freshwater habitat has been adopted independently a number of times 
by some ancestral marine genus such as Lironeca. Monod (1931) also mentions the 
above points, but despite this the name Ichthyoxenus still seems to survive for 
certain freshwater species. More recently, Fryer (1965) adopts the view that the 
freshwater species have been derived from some common marine ancestor, and that 
morphologically they are indistinguishable from the widespread marine genus 
Lironeca. Fryer thus refers expansus to Lironeca, and considers Ichthyoxenus of 
Herklots 1870 to be a junior synonym of Lironeca Leach 1818. 

Two further species are described by Fryer from Lake Tanganyika, Lironeca 
tanganyikae Fryer 1965 and Lironeca enigmatica Fryer 1968. The former species, 
tanganyikae, has been recorded only from the mouth cavity of the littoral cichlid 
fish Simochromis diagramma (Giinther), while enigmatica was taken from the body 
and gill cavity of the clupeids Limnothrissa miodon (Boulenger) and Stolonothrissa 
tanganicae Regan. 

An examination of a collection of the cichlid fishes Lamprologus elongatus 
Boulenger and Lamprologus pleurostigma Boulenger from Lake Tanganyika has 
provided a number of isopod parasites, with both mature male and female stages 
present. These specimens represent a hitherto undescribed species which I name 
Lironeca africana. 

Bull. Br. Mus. nat. Hist. (Zool.) 21, 8 



330 R. J. LINCOLN 

Lironeca africana sp. nov. 
(Text-figs la-g, 2a-j, 3a-q) 

MATERIAL EXAMINED. Holotype female, length u-o mm, ovigerous: allotype 
male found in association with holotype, length 5-0 mm: paratypes, 5, $$, length 
7 -0-14 -o mm, width 3 -5-5 -o mm ; 4, <$$, length 4 -0-5 -5 mm, width i -0-2 -o mm. All 
specimens deposited in the British Museum (Natural History), accession numbers; 
holotype, 1970 : 438; allotype, 1970 : 439; paratypes, 1970 : 440. 

FEMALE, (holotype). Body size, length n-o mm, width 4-5 mm, height including 
the marsupium 4-0 mm. Shape of body in dorsal, ventral and lateral view given in 
Figs la, 2a, ib respectively. Head, ratio of length to maximum width about 
i : 1-5, shape triangular, broadly rounded at the front and curved slightly down- 
wards towards the base of the antennae ; eyes large, laterally placed ; head somewhat 
sunken into the peraeon ; chromatophores evenly distributed on dorsal side of head. 
Peraeon tergite i about i| times the length of tergite 2, tergites 2-6 subequal, 
tergite 7 only half length of tergite 6; peraeon tergite i, antero-lateral projection 
reaching forward to mid-eye level, anterior and posterior margins broadly rounded ; 
peraeon tergite i almost flat dorsally, tergites 2-7 becoming progressively more 
convex ; maximum width of peraeon is reached by coxal plates of segment 4 ; coxal 
plates reach progressively nearer to posterior margin of tergites from 1-7 ; posterior 
margin of peraeon tergite 7 broadly rounded. Pleon immersed a little in peraeon 
tergite 7, all segments free laterally, length of segments increasing slightly from 1-5 ; pos- 
terior margin of pleon segment 4, and especially 5, sinuous. Telson, length twice width, 
broadly rounded, with ridge running from antero-lateral corners towards the centre. 

Appendages; antenna i and 2 each with 7 segments, slender, rounded and only 
just reaching beyond forward projection of peraeon tergite i, antenna i slightly 
longer than antenna 2 (Fig. 2b, c) ; mandible (Fig. 2d) with slender inscisor process 
pointed at tip, molar process rounded, palp 3-segmented with 2-3 apical setae; 
maxilla i (Fig. 2e) slender, carrying 4 recurved spines on distal tip ; maxilla 2 (Fig. 2f) 
small, bilobed, each lobe bearing a pair of hooked spines; maxilliped (Fig. 2g) well 
developed, broad, with a 2 segmented palp terminating in 4 strongly recurved spines. 
Peraeopods 1-7 of similar shape (Fig. ic, d), basis and ischium strongly developed in 
anterior peraeopods, less so in posterior peraeopods, merus and carpus very short, 
propodus prominent, dactylus strongly curved and pointed, apex of dactylus fitting 
into a carpal groove, very well formed articulation between dactylus and propodus, 
no spines present on the peraeopods. Pleopods fleshy, leaf-like, without marginal 
setae (Fig. ic, f) ; uropods (Fig. ig), inner ramus just a little longer than outer, both 
rami apically rounded, projecting fractionally beyond margin of telson. 

MALE, (allotype) found associated with holotype. The male is very much smaller 
than the female, length 5 mm, width 2 mm ; general form of the body and appendages 
resemble the female, and reference will therefore be made only to the main points of 
difference. Shape of body in dorsal view as in Fig. 2h. Head with rounded eye 
lobes, produced forward into a short, somewhat square-ended rostrum, only weakly 
immersed into peraeon; peraeon slightly asymmetrical, tergite i is i| times length 



LIRONECA (ISOPODA; CYMOTHOIDAE) 





FIG. i. Lironeca africana sp. nov. $ Holotype: a, body entire, dorsal view; b, body entire, 
lateral; c, peraeopod i left, ventral view; d, peraeopod 7 left, ventral view; e, pleopod i 
left, ventral view; f, pleopod 2 left, ventral view; g, uropod left, ventral view. Bar scale 
i mm. 



332 



R. J. LINCOLN 




FIG. 2. Lironeca africana sp. nov. a-g $ Holotype: a, body entire, ventral; b, antenna i ; 
c, antenna 2; d, mandible, e, maxilla i; f, maxilla 2; g, maxilliped; h-j <$ allotype: 
h, body entire, dorsal; i, pleopod i, left; j, pleopod 2, left. Bar scale i mm. 



LIRONECA (ISOPODA; CYMOTHOIDAE) 333 

of tergites 2-7, maximum width at the level of segment 3, posterior margin of tergite 
7 deeply rounded ; paired penis lobes present on ventral surface of peraeon segment 7 ; 
chromatophores on both dorsal and ventral surface of peraeon; body less convex 
than the female. Pleon segment i immersed laterally in peraeon, segments 2-5 
free; telson broad and flat, length about half of width, posterior corners round 
becoming straight in the centre. 

Appendages; antenna i (Fig. 2h) rounded in cross section, 7 segmented; antenna 
2 much narrower and consisting of 8 segments; mandible, maxilla i, maxilla 2 and 
maxilliped as in female except the apical hooks are much less pronounced; pleopod 
2, endopod, bearing appendix masculina (Fig. 2J), pleopod i, fleshy and leaf -like 
(Fig. 2i). 

First marsupial stage. This is mentioned only briefly as it is the structure of the 
second marsupial stage which is of greater interest. Body shape as in Fig. 3m; 
head not immersed in peraeon; peraeon tergite i with markedly sinuous posterior 
margin, segment 7 reduced in size and continuous with pleon; telson long and 
rounded, no marginal setae. 

Appendages; antenna i consists of 7 segments, much dilated, without aesthetascs; 
antenna 2 of 8 segments, slender, slightly longer than antenna i. Peraeopods 1-6 
of similar size (Fig. 3n, o), no hooks or spines present. Pleopods 1-5 fleshy, no 
marginal setae; uropods projecting well beyond extremity of telson, no marginal 
setae or terminal spine (Fig. 3p, q). 

Second marsupial stage. Body, length about 3 mm, width about i mm ; 65 young 
specimens taken from the marsupium of a single female ; shape in dorsal view as in 
Fig. 3a. Head with rounded eye lobes and forwardly produced rostrum as seen in 
the male ; eyes large. Peraeon tergite i almost twice length of tergites 2-6, anterior 
and posterior margin of tergite i broadly rounded, segment 7 reduced and con- 
tinuous with pleon, dorsal side of peraeon flattened; no penis lobes evident. Pleon 
rectangular, all segments free laterally; telson slightly longer than wide, rounded 
posterior margin bearing about 13 long plumose setae. 

Appendages ; antenna i (Fig. 3b) rounded in cross section and dilated, 7 segments, 
of which 4-7 carry prominent groups of aesthetascs; antenna 2 (Fig. 3b) slender, a 
little longer than antenna i, 8 segmented with a small group of terminal setae; 
mandible, maxilla i, maxilla 2, and maxilliped similar to those of female except 
much smaller and the terminal hooks evident only as small rounded tubercles. 
Peraeopods increasing in length very slightly from 1-7, basis and ischium well 
developed, merus and carpus short, propodus strong, and dactylus long, pointed, 
and sharply curved; peraeopods 1-3, merus with long spine-like seta on antero- 
lateral corner (Fig. 3c), also a characteristic number of hooks on inner side of each 
peraeopod, peraeopod 1-2 with a pair of hooks on propodus (Fig. 30), peraeopods 
3-4 have a pair of large hooks on propodus and a single curved spine on carpus 
(Fig. 3d), peraeopod 5 with two propodal and two carpal spines (Fig. 36), peraeopod 
6 bears two spines on propodus, three on carpus, and a single one on the merus 
(Fig. 31). Pleopods bilobed (Figs 3g-k), leaf -like with long marginal plumose setae, 
setae absent from endopods of pleopods 3-5 ; peduncle of pleopods 1-5 each with a 



334 



R. J. LINCOLN 




FIG. 3. Lironeca africana sp. nov. a-1, second marsupial stage: a, body entire, dorsal; 
b, antennae i & 2 ; c, peraeopod i, left; d, peraeopod, 3, left; e, peraeopod 5, left; f, perae- 
opod 6, left; g, pleopod i, left; h, pleopod 2, left; i, pleopod 3, left; j, pleopod 4, left; 
k, pleopod 5, left; 1, uropod & telson entire, m-q, first marsupial stage; m, body entire, 
dorsal; n, peraeopod i, left; o, peraeopod 6, left; p, pleopod i, left; q, pleopod 2, left; 
bar scale 0-5 mm. 



LIRONECA (ISOPODA; CYMOTHOIDAE) 335 

set of coupling spines ; exact number of marginal setae on the pleopods is somewhat 
variable; uropods (Fig. 3!) with long plumose setae, the exopod bearing a single 
large curved spine at the distal tip. 

REMARKS. As might be expected in a group which shows marked polymorphism, 
there are some small morphological variations in the paratype material. It is 
important to appreciate this variability when attempting to identify material such 
as this, and the lack of such appreciation has undoubtedly been partly responsible 
for the very large number of species which have been described in the past. 

The shape of the telson in females shows some variation from that of the holotype, 
and may be broad, almost rectangular, with a stright posterior margin. The 
peraeon in one of the specimens is asymmetrical, and there are some differences in 
proportional lengths of the peraeon tergites. The first tergite of the peraeon is not 
always markedly longer than tergites 2-6, but in all specimens the last tergite is 
reduced in length. In the holotype the peraeopods are all approximately the same 
length, while in the largest female paratype there is a small increase in the length of 
the peraeopods from 1-7, reflected in the size of the ischial segment, the length of 
the other segments remaining unchanged. In the males it is important to note that 
the body may show signs of asymmetry, and there is a wide range of body size, some 
specimens being very small. 

The parasites were all taken from the mouth cavities of the cichlid fishes Lampro- 
logus elongatus Boulenger and Lamprologus pleurostigma Boulenger. The fishes 
examined were part of an extensive collection taken from Lake Tanganyika and 
deposited in the collections of the British Museum (Natural History). The female 
parasite was attached to the tongue of the fish, with the long dactylae of the peraeo- 
pods embedded deeply into the tissues of the host. The isopod was situated with its 
head innermost, the posterior end of the body being about level with the angle of the 
host's jaw. Despite the large size of the parasite the fish was still able to close its 
mouth fully. The male parasite was found in close association with the female, 
attached either to the side of the mouth or the inside of the gill chamber. Only one 
male was found in the presence of each female. 

Lironeca africana is readily distinguished from other freshwater African species 
by the general outline of the body and the contour of the posterior margin of peraeon 
tergite 7. Lironeca expansus is broadly elliptical in outline and the posterior margin 
of tergite 7 has a deeply concave contour, with the pleon strongly immersed into the 
peraeon. The head is also deeply set into the peraeon and projects very little, if 
any, beyond the general outline of the body. Lironeca tanganyikae has an oval out- 
line, somewhat similar to africana, but the margin of tergite 7 is broadly convex with 
the median part straight. The pleon is only moderately immersed into the peraeon. 
In Lironeca africana the body is more elongate, the convex posterior margin of tergite 
7 is an even curve, and the pleon is but weakly immersed. The characters men- 
tioned above refer to the mature female stage. 

The isopods belonging to the family Cymothoidae are protandrous hermaphrodites 
and it has been usual practice to base the description of a new species on the adult 
female stage. In the case of Lironeca enigmatica, described by Fryer from Lake 



336 R. J. LINCOLN 

Tanganyika, the fully mature female was not available and the type series consists 
of a number of young male specimens varying both in size and in degree of develop- 
ment. The presence of setae on the pleopods and uropods indicates that these 
isopods were capable of free swimming, and it still remains to be shown whether or 
not the adult parasite is to be associated with the same species of fishes from which 
the enigmatica material was taken. Bearing this in mind, and appreciating the 
marked transformation which takes place when the parasite takes up permanent 
residence with the host, it could prove particularly difficult to establish the true 
identity of the adult female. To separate enigmatica and africana it is pointless to 
look at the male or female of the latter as they are no longer free swimming, and 
have lost all traces of pleopodal and uropodal setae. It only remains to compare 
enigmatica with the late marsupial stage of africana, which does have a full setal 
compliment in preparation for its early free swimming existence. These can easily 
be distinguished on the basis of the number of spines and hooks on the peraeopods 
and also the shape of the telson. 

A close investigation of the characters possessed by the late marsupial juvenile 
stage of cymothoids may eventually provide an answer to some of the many problems 
which face the taxonomist working with this group. At the present time there are 
many genera which are separated on quite unreliable features and, as Fryer points 
out, it is quite possible to refer a given specimen to any of a number of genera. This 
has led to some genera receiving a large number of species, Lironeca being a good 
example, while others have remained monotypic. 



The discovery of this parasite in association with cichlid fishes of the genus 
Lamprologus revives the question of the origin of the Lake Tanganyika isopod fauna. 
The cymothoids found in the Lake have very obvious affinities with marine members 
of the group, with which they no doubt share a common marine ancestral group. 
The problem is to establish when the isopods made their entry into the Lake and 
acquired freshwater status. Three main possibilities exist, although the first of 
these to be outlined can be said to have little support from present knowledge of the 
geology and biology of the Lakes region of eastern Africa. In a book 'The Tan- 
ganyika Problem' published in 1903, Moore discussed the apparent affinities of the 
Lake fauna, with special reference to the molluscan groups, and arrived at the con- 
clusion that the fauna had evolved from an ancient Jurassic marine stock which 
became isolated when the Lake was cut off from the sea. If the fauna originated in 
in this way, the transition of the animals to a freshwater existence would have 
occurred within the Lake, and the parasites could have made the change in associa- 
tion with their fish hosts. But, as has been noted above this theory of the origin 
of the Lake fauna and of the Lake itself has now been largely abandoned. 

From the evidence of the present distribution of the freshwater parasites it seems 
most likely that the isopods entered the Lake from a connection with the Congo 
river system. Lake Tanganyika is the only one of the Lakes in eastern Africa which 
is known to have the parasites, and the Congo River system is the only one of the 
rivers known to have a freshwater cymothoid fauna. If the isopods entered the 



LIRONECA (ISOPODA; CYMOTHOIDAE) 337 

Lake region at a time before the isolation of Lake Tanganyika itself, one would 
expect to find them in some of the other bodies of freshwater, but this is not the case. 
It is now generally accepted that Tanganyika was formed sometime during the 
Pleistocene, but the question remains whether the isopods reached the Lake soon 
after its isolation, or whether they moved in from the Congo through some river 
connection in comparatively recent times. The evidence, limited as it is, tends to 
support the former possibility. 

The two species of parasite found in the Lake for which a final host is known, 
tanganyikae and africana, are both associated with cichlid fishes, and show strict 
host specificity. This in itself suggests a long association, and the cichlid fishes 
represent a rich endemic element in the Tanganyika fauna. The third cymothoid 
known from the Lake, enigmatica, was taken from the clupeids Stolonothrissa and 
Limnothrissa, but these isopods were still juvenile and capable of free swimming, 
and the identity of their final host has not yet been established. Fryer (1968) points 
out that these clupeids have marine ancestors and the parasite may have made the 
transition to freshwater with the fish, and further suggests that this could represent 
a separate invasion of the Lake by the isopods. This cannot be disproved, but it 
is worth noting that the fishes in question are pelagic species which aggregate in 
large shoals, and are sufficiently abundant to be commercially exploited (Coulter 
1970). Fish which move freely about the Lake and undergo marked diurnal migra- 
tion would provide ideal dispersal for the juvenile free living stages of a parasite, 
and it seems quite probable that they are acting only as temporary intermediate 
hosts for the isopods. The fish are preyed upon by numerous carnivorous fish 
species, including some cichlids, and this may be the step by which the parasite 
reaches its final host. The speculation will remain until the adult enigmatica has 
been found, but if the view outlined above is found to be true, and the adult parasite 
is associated with a cichlid fish, it will be less of an enigma than the name was 
intended to suggest. 

The cymothoid, expansus, found widely in the waters of the Congo, is without 
doubt a very close ally of the Tanganyika species. It too would seem to be host 
specific, on Eugnathichthys eetweldii, as I can find no record of it having been taken 
from any other fish, not even from a closely related species of the same citharinid 
fish genus which is also common in the Congo river. 

The obvious affinities of the freshwater cymothoids in Africa one with another, 
and their very restricted distribution, indicates an entry into Lake Tanganyika from 
the Congo river system. The strict host specificity of the parasites and the endemi- 
city of the host fishes points to an early invasion of the Lake rather than an entry in 
more recent times. However, the nature of this brief summary must remain 
speculative until a great deal more is known of the isopods of the African continent. 



338 R. J. LINCOLN 

REFERENCES 
BRIAN, A. & DARTEVELLE, E. 1949. Contribution a l'6tude des Isopodes marins et fluviatiles du 

Congo. Annls Mus. r. Congo Beige. C, Zoologie 1 (2) : 77-208. 

COULTER, G. W. 1970. Population changes within a group of fish species in Lake Tangan- 
yika following their exploitation. /. Fish Biol. 2 : 329-353. 
DARTEVELLE, E. 1939. Ichtyoxenus expansus, Isopode parasite dulcaquicole. Rev. Zool. 

Bot. afr. 33 (i) : 16-17. 
FRYER, G. 1965. A new isopod of the genus Lironeca, parasitic on the cichlid fish of Lake 

Tanganyika. Rev. Zool. Bot. afr. 71 (3-4) : 376-384. 
- 1968. A new parasitic isopod of the family Cymothoidae from clupeid fishes of Lake 

Tanganyika a further Lake Tanganyika enigma. /. Zool., Lond. 156 : 35-43. 
GROSSE, J. P. 1963. Le milieu aquatique et Ideologic des poissons dans la region de Yan- 

gambi. Annls Mus. r. Afr. cent., Sci. Zool. 116 : 113-271. 
MIERS, E. J. 1880. On a collection of Crustacea from the Malaysian region. IV. Penaeidae, 

Stomatopoda, Isopoda, Suctoria and Xiphosura. Ann. Mag. Nat. Hist. 5 : 457-472. 
MONOD, T. 1931. Sur quelques Crustaces aquatiques d'Afrique (Cameroun et Congo). Rev. 

Zool. Bot. afr. 21 (i) : 1-36. 

MOORE, J. E. S. 1903. The Tanganyika Problem. London. 
NAME, W. G. VAN. 1920. Isopods collected by the American Museum Congo Expedition. 

Bull. Am. Mus. not. Hist. 43 : 41-108. 



Dr. R. J. LINCOLN 

Department of Zoology 

BRITISH MUSEUM (NATURAL HISTORY) 

CROMWELL ROAD 

LONDON, S.W-7 



EUNICE MANIHINE SP. NOV. (POLYCHAETA: 

EUNICIDAE), A MEMBER OF THE FLAVUS- 

BIDENTATE GROUP FROM THE WESTERN 

EQUATORIAL INDIAN OCEAN 

By M. R. LONGBOTTOMt 



SYNOPSIS 

A new species of the genus Eunice Cuvier, 1817, is described from 421 m depth in the western 
equatorial Indian Ocean. The characteristic features are: yellow, bidentate acicular setae, up 
to 4 per parapodium; up to 6 acicula per parapodium; branchiae, with few filaments, present 
from setiger 8 to 43; smooth occipital tentacles, the median reaching to setiger 24. 

INTRODUCTION 

THE few surveys of polychaetes carried out in the western equatorial Indian Ocean 
have been restricted to intertidal and shallow-water habitats, rarely exceeding 50 m 
depth (Crossland, 1904), and little is known of the fauna of the rough bottoms at 
greater depths. During the Royal Society Indian Ocean Deep Slope Fishing 
Expedition (January/February, 1969), vertical bottom lines were fished from the 
FRV Manihine around the islands and banks in this region of the Indian Ocean 
(Forster et al., 1970). One of these lines foul-hooked a piece of coral debris at 421 m, 
in which was a eunicid worm ; further examination showed this to be an undescribed 
species of Eunice. 

The genus Eunice Cuvier, 1817, consists of a large number of valid species (approxi- 
mately 140) for which the common specific characters and their variations are 
discussed in a recent review (Fauchald, 1970). Briefly, these are the form and 
colour of the acicular setae (subacicular hooks), the distribution and form of the 
branchiae, and the length and form of the occipital tentacles. Combinations of 
these and, frequently, other characters serve to distinguish the species, only rarely is 
one of these major characters alone sufficient. Major divisions, based on the form 
and colour of the acicular setae, were suggested by Hartman (1944), resulting in 
four groups of those species for which the relevant details were known. This 
system has been expanded to include a greater number of valid species, and, based 
on the distribution of the branchiae, five subdivisions have been proposed for each 
of seven groups (Fauchald, 1970). 

DESCRIPTION 

The single specimen of Eunice manihine sp. nov. consists of 127 setigers and is 
approximately 75 mm long (9 mm from the tip of the palps to the posterior edge of 
the tenth setiger on the dorsal side, following Fauchald, 1970) and 5 mm wide. The 

t Dr. Longbottom died August 6, 1971 
Bull. BY. Mus. nat. Hist. (Zool.) 21, 8 



34 



M. R. LONGBOTTOM 



specimen is cylindrical in the anterior region, and becomes flattened towards the 
posterior. 

The anterior margin of the prostomium is distinctly notched between the palps 
(Fig. i). The length of the prostomium is slightly greater than half the breadth. 
The prostomium bears a pair of eyes posterior to the gap between the bases of the 




5mm 




FIGS 1-7. Eunice manihine. i. Anterior end, dorsal view (left outer occipital tentacle 
missing). 2. Setigers 14-17 right side, dorsal view. 3. Posterior end, left lateral 
and slightly ventral view. 4. Maxillary plates, dorsal view. 5. Setigers 7-10, dorsal 
view. 6. Setigers 7-10, ventral view. 7. Setigers 7-10, diagrammatic representa- 
tion, dorsal side firm lines. 



EUNICE MANIHINE 341 

inner and outer occipital tentacles (Fig. i). Five smooth and very long, slender 
tentacles are present ; the unpaired median tentacle reaching back to setiger 24, the 
inner lateral tentacles reaching setiger 12 and the outer laterals setiger 3. The 
first peristomial segment is two thirds the length of the prostomium; the second 
peristomial segment is half the length of the first and the same length as the first 
setiger. The peristomial cirri are smooth and reach just beyond the posterior border 
of the prostomium. 

The maxillae are well developed (Fig. 4); maxilla I is falcate; maxilla II has 12 
teeth left and n teeth right; maxilla III has 12 teeth left; left maxilla IV has n 
teeth and the combined right maxillae III +IV have 13 teeth ; each maxilla V has 
i tooth. 







1mm 






n 



15 



16 17 



12 



FIGS 8-17. Eunice manihine. 8-n. Parapodia from right side, anterior view. 8. 
Setiger 4. 9. Setiger 34. 10. Setiger 60. n. Setiger 90. 12. Capillary seta from 
setiger 90. 13. Comb seta from setiger 90. 14. Falcigerous seta from setiger 34. 
15. Acicular seta (subacicular hook) from setiger 34. 16. Aciculum from setiger 34. 
17. Aciculum from setiger 60. 



342 M. R. LONGBOTTOM 

Branchiae are present only in the anterior third of the specimen: from setiger 8 
to setiger 43. The branchiae consist of a single filament anteriorly (Fig. i) with a 
maximum of 3 filaments between setigers 17 and 22 (Fig. 2), decreasing posteriorly 
to a single filament. The filaments are thinner than the dorsal cirri in the branchial 
region (Fig. 2). The dorsal cirri are smooth, anteriorly with a stout base and 
elongated tip (Fig. 8), posteriorly digitiform (Figs 3 & n). The ventral cirri are 
subulate (Figs 3, 6, 8-n). Two pairs of anal cirri are present (Fig. 3), the dorsal 
pair long and the ventral pair very short. 

In the majority of the parapodia there are, superiorly, between 10-18 long, slightly 
limbate, capillary setae with weak striations (Fig. 12), and, towards the posterior, 
5-9 comb setae with 14-16 teeth, usually with the outer tooth on one side appreciably 
longer (Fig. 13). Inferiorly, there are 7-9 falcigerous setae, the blades of which are 
strongly bidentate, with a rounded hood projecting just beyond the tip (Fig. 14). 
The head of the shaft is slightly enlarged. The acicula are yellow with slightly 
curved tips (Figs 16 & 17). An average of 3 or 4 are present in each parapodium, 
with up to 6 in the middle setigers, decreasing to 2 posteriorly. The inferiorly 
situated acicular setae (subacicular hooks) are first present, singly, from setiger 
27/28, with 3-4 per parapodium from setiger 31 throughout the remainder of the 
setigers. The acicular setae are yellow, bidentate, and hooded (Fig. 15). 

DISTRIBUTION 

E. manihine is known from one specimen found in a piece of coral debris foul- 
hooked on a bottom fishing line from 421 m depth, south of Menai Is, Cosmoledo 
(10 S, 47 E), in the western equatorial Indian Ocean. 

The type specimen (Ref. no. ZB 1971-1) which is preserved in alcohol has been 
deposited in the British Museum (Natural History). 

DISCUSSION 

No single character is sufficient by itself to distinguish E. manihine from other 
members of the genus; the separation is based on a combination of the following 
characters: yellow, bidentate, acicular setae; branchiae of few filaments restricted 
to the anterior third of the body; very long, smooth, occipital tentacles; the form of 
the maxillary plates; and the large number of acicula per parapodium. The first 
two characters are common to many species of Eunice and are discussed in greater 
detail beiow. References in the literature to occipital tentacles reaching to near or 
beyond setiger 20 are uncommon : E. antillensis has been recorded with the median 
tentacle to setiger 20 (Ehlers, 1887), E. auriculata to setiger 18 (Treadwell, 1901), 
E. palauensis to setiger 18 (Okuda, 1937), and E. tibiana (Pourtales, 1863) to setiger 
22 (Izuka, 1912). Of these species only E. antillensis is a member of the flavus- 
bidentate group. However, the degree of variation within a species is not known, 
and to some extent is dependent upon the state of preservation. This also applies 
to the form of the tentacles, whether they are wrinkled or clearly articulated. The 
presence of as many as 6 acicula per parapodium is very unusual, contrasting with 
the more normal number of 1-3 for species of Eunice. The number of teeth on the 
maxillary plate is also greater than is usually found in the genus. 



EUNICE MANIHINE 343 

Adoption of the system of subdivision proposed by Hartman (1944) and Fauchald 
(1970) enables comparisons to be made more easily within this large genus. E. 
manihine, in possessing yellow, bidentate acicular setae, clearly belongs to the 
flavus-bidentate (A) group of Hartman (1944), and from the distribution of the 
branchiae (commencing before setiger 10 and not present after setiger 100), to 
subdivision i of Fauchald (1970). As the numbers of setigers may vary in worms 
of different sizes, it is suggested that the definition of this subdivision should be 
expanded to place the emphasis on the branchiae being present in the anterior third 
of the body rather than within a definite number of setigers irrespective of the size 
of the worm. 

Fourteen species are listed by Fauchald (1970) within this group Ai to which 
should be added E. semisegregata Fauchald, 1970, and from which should be 
removed E. benedicti (Verrill, 1885) a synonym of E. pennata (Miiller, 1776) (see 
Pettibone, 1963) and E. norvegica (Linnaeus, 1767). The latter, in having black 
bidentate acicular setae, and branchiae present from setiger 3 to within a few setigers 
of the anus (Pettibone, 1963), should be placed in the fuscus-bidentate (B) group, 
subdivision 2. All the previously described species in the Ai grouping have the 
branchiae commencing on setiger 3. Fauchald (1970) has suggested that this is a 
constant feature within a species, though small variations have been recorded, as for 
example in E. pennata (Pettibone, 1963). Thus, E. manihine is the sole member of 
the flavus-bidentate group in which the branchiae commence before setiger 10, but 
after setiger 6. Apart from E. manihine, only two species E. megabranchia 
Fauchald, 1970 and E. validobranchiata Monro, 1937 in the Ai group have smooth 
occipital tentacles, the remainder have articulated or moniliform tentacles. E. 
megabranchia (from 894 m depth in the Gulf of California) and E. validobranchiata 
(from 1046 m depth off the South Arabian coast) differ only in the length and shape 
of the guards of the falcigerous setae and the form of the ventral cirri; the other 
characters are very similar and for comparison with E. manihine may be considered 
identical. Specimens of these two species of a similar size to the E. manihine 
specimen differ in having branchiae with large numbers of filaments (up to 45) 
commencing on setiger 3, appreciably shorter occipital tentacles, fewer acicular 
setae per parapodium and fewer teeth on the maxillary plates. 

Of the eunicid polychaetes reported from East Africa (Crossland, 1904), only one 
species of the flavus-bidentate group was reported E. tubifex Crossland, 1904. 
This belongs to Fauchald's subdivision 4, in which branchiae are present from after 
setiger 10 to the posterior. 

The type specimen of E. manihine shows an abnormality in the segmentation of 
setigers 8 and 9 (Figs 5 & 6), resulting in a short spiral (Fig. 7) similar to the form 
Buchanan (1893) found in specimens of Lumbriconereis impatiens (Lumbrinereis 
tetraura}. It was not possible to examine the internal anatomy of the single speci- 
men available. 

ACKNOWLEDGEMENTS 

I am most grateful to Dr. O. Hartman and Professor J. H. Day for confirming the 
separate identity of the species, and to Dr. J. D. George and Mr. P. M. David for 



344 M - R - LONGBOTTOM 

reading the manuscript. Study leave was granted by the University of Newcastle 
upon Tyne for the period of the expedition, which was financed by the Royal Society. 

REFERENCES 

BUCHANAN, F. 1893. Peculiarities in the segmentation of certain polychaetes. Q. Jl 

microsc. Sci. 34 : 529-544. 
CROSSLAND, C. 1904. The marine fauna of Zanzibar and British East Africa, from collections 

made by Cyril Grassland in the years 1901 and 1902. The Polychaeta. Part III. With 

which is incorporated the account of Stanley Gardiner's collection made in the Maldive 

Archipelago in the year 1899. Proc. zool. Soc. Land. 1 : 287-330. 
EHLERS, E. 1887. Report on "the annelids of the dredging expedition of the U.S. Coast Survey 

steamer Blake. Mem. Mus. comp. Zool. Harv. 15 : 1-335. 
FAUCHALD, K. 1970. Polychaetous annelids of the families Eunicidae, Lumbrineridae, 

Iphitimidae, Arabellidae, Lysaretidae and Dorvilleidae from Western Mexico. Allan 

Hancock Monographs in Marine Biology 5 : 1-335. 
FORSTER, G. R., BADCOCK, J. R., LONGBOTTOM, M. R., MERRETT, N. R. & THOMSON, K. S. 

1970. Results of the Royal Society Indian Ocean Deep Slope Fishing Expedition, 1969 

Proc. R. Soc. B. 175 : 367-404. 
HARTMAN, O. 1944. Polychaetous annelids. Part 5. Eunicea. Allan Hancock Pacific 

Exped. 10 : 1-238. 
IZUKA, A. 1912. The errantiate Polychaeta of Japan. /. Coll. Sci. imp. Univ. Tokyo 30 (2) : 

1-262. 
OKUDA, S. 1937. Polychaetous annelids from the Palau Islands and adjacent waters, the 

South Sea Islands. Bull, biogeogr. Soc. Japan 7 : 257-316. 
PETTIBONE, M. H. 1963. Marine polychaete worms of the New England region. I. Families 

Aphroditidae through Trochochaetidae. Bull. U.S. natn. Mus. 227 : 1-356. 
TREADWELL, A. L. 1901. The polychaetous annelids of Porto Rico. Bull. U.S. Fish Commn 

20 : 181-210. 



Dr. M. R. LONGBOTTOM 

NATIONAL INSTITUTE OF OCEANOGRAPHY 

WORMLEY 

GODALMING, SURREY 



ORIGINAL DESCRIPTION OF THE GANGETIC 

DOLPHNI, PLATANISTA GANGETIC A, 
ATTRIBUTED TO WILLIAM ROXBURGH 

By G. PILLERI 

IN cetological literature the original description of the Gangetic dolphin is usually 
attributed to Heinrich Julius Lebeck. In the third volume of "Neue Schriften der 
Gesellschaft Naturforschender Freunde zu Berlin" that appeared in 1801, this 
author published a short paper entitled "Delphinus gangeticus beschrieben von 
Herrn Heinrich Julius Lebeck zu Trankenbar". "Trankenbar" obviously corres- 
ponds to the locality "Tranquebar" in southeast Madras. 

In the same year, 1801, Volume 7 of "Asiatick Researches" was published in 
Calcutta and contained a paper of William Roxburgh entitled "An Account of a 
new Species of Delphinus, an Inhabitant of the Ganges". 

It is, therefore, virtually certain that the two descriptions of the Gangetic dolphin 
were published in the same year, but the reviews contain no definite indication as 
to which of the two appeared first. 

Roxburgh's paper was republished in 1803 by Maiden and Wilson of London in a 
second edition of "Asiatick Researches, printed verbatim from the Calcutta edition". 

In recent reports (e.g. Hershkovitz 1966), data taken from literature onPlatanista 
gangetica are frequently referred back to the paper published by Roxburgh in 1803, 
the date of publication of the London edition. 

When G. Cuvier (1823) changed the designation of the genus Delphinus to Platan- 
ista, he claimed priority for Lebeck. E. Home (1818), on the other hand, in his 
paper on the morphology of the teeth of the Gangetic dolphin, referred to Roxburgh 
as the original author. M. F. Cuvier (1836) who, in nearly all other respects, based 
his work on the account of his brother (G. Cuvier 1823), expressed a doubt as to 
whom to attribute the original work. Referring to the paper of Home, he added a 
footnote at the bottom of page 252 that reads as follows: "T.VII, p. 170, pi. Ill 
(e"dit. de Londres, 1803, in-4). M. Home dans sa description des dents de ce 
dauphin, cite le memoire de Roxburgh comme ayant paru en 1721 dans les me"moires 
de la societe de Calcutta. Ces memoires paraissaient-ils alors? L'e"dition de 
Londres est une copie de celle de Calcutta". "1721" is obviously a printing error. 

When John Anderson (1833-1900) first took up his activities in India he was 
staying with his brother, a doctor and botanist, in the house built in the Botanical 
Gardens by Roxburgh. On page 418 of the monograph he published in 1878 on 
Platanista and Orcaella, Anderson mentioned that Roxburgh claimed to have 
published the first description of the Gangetic dolphin, but made no further comment 
on the fact. 

A comparison of the two papers of Lebeck and Roxburgh reveals a striking 

Bull. Br. Mus. nat. Hist. (Zool.) 21, 8 



346 G. PILLERI 

similitude in the text, both as regards their presentation and the zoological data they 
contain. Here, I should like to give a brief summary of the lives of the two authors. 

William Roxburgh. Born at Underwood, Craigie, Ayrshire, on June 3rd, 1751, 
William Roxburgh became a distinguished botanist. He studied in Edinburgh 
under John Cope and it was Cope who procured him an engagement as assistant 
surgeon on a ship of the East Indian Company. After several voyages to the 
tropics, he was granted a degree of Doctor of Medicine in Edinburgh and from 1776- 
1778 worked in the General Hospital in Madras. In 1780, he became full surgeon 
and a year later took up a post at Cocanada, in the delta of the Godavery River, 
where he was mainly occupied with applied botany. It was during this period that 
he undertook his very extensive collection of plants and illustrations of the Indian 
flora. In 1793 he was appointed Superintendent of the Botanical Gardens of 
Calcutta but four years later he was forced to return to Scotland on account of ill 
health. He went back to Calcutta in 1799, but had to leave India again in 1805. 
His state of health continued to deteriorate and he sought recovery at the Cape and 
in St. Helena, but in vain. He finally returned to Scotland and died in Edinburgh 
on February i5th, 1815. 

The numerous and very comprehensive studies undertaken by William Roxburgh, 
some of which were published after his death by Wallich, constitute a valuable 
contribution to the knowledge of the Indian, and more particularly, of the Bengalese 
flora (see Lee 1897). He also compiled reports on water insects, silk worms and land 
winds. The work on the Gangetic dolphin (1801) falls in a period of Roxburgh's 
life already overshadowed by the outbreak of his serious illness. 

Heinrich Julius Lebeck. In spite of extensive investigations in a large number of 
libraries at home and abroad, I have not been able to uncover any biographical 
details on the life of Lebeck. The only reference I found to his life was contained in 
the book of G. Cuvier (1823) wno mentioned that he was a "Danish missionary in 
Tranquebar". However, this piece of information is obviously inaccurate as Gosch 
(1875), in his book on early Danish zoological literature, describes Lebeck as a 
German missionary. Furthermore, the biographical lexicon in the royal library in 
Copenhagen contains no mention of Lebeck. 

The descriptions of Lebeck and Roxburgh 

It should be pointed out that Roxburgh stresses the discovery of a new species in 
the title of his work, "An Account of a New Species of Delphinus", and Lebeck does 
not. Both descriptions refer to a male specimen from the Calcutta region of exactly 
the same size (!). What is even more significant is that not only were the measure- 
ments quoted in the two papers identical, but in Lebeck's paper British linear 
measurements and British avoirdupois weights were used. If the original contribu- 
tion had been German one would logically expect the author to have employed 
German units. The two reports start with Linnaeus and state that the Gangetic 
dolphin is a fifth, new species to be added to the four species of dolphins described 
by Linnaeus. The head, teeth, tongue, eyes, external auditory passages, fins and 
genitals are described in the same order. Both authors found nematodes and plant 
seeds in the stomach. Both mention the thickness and strawyellow colouring of the 



THE GANGETIC DOLPHIN PLATANISTA GANGETICA 347 

blubber as well as the fact that dolphins oils are used for medicinal purposes by the 
natives. 

Not only is the order in which the organs are described in the diagnosis of the 
species identical, but also, whole sentences are repeated word for word. 

A comparison of the texts leads to the conclusion that one of the two authors was 
acquainted with the text of the other before he published his own work. It is also 
significant that Lebeck mentions a "zweiten mir gezeigten Russel eines solchen 
Tieres" 1 (page 282). I presume that this rostrum could only have been shown to 
him within the natural science circles of Calcutta, the town where research on the 
Gangetic dolphin originated with the collections and observations of Roxburgh, 
Blyth and, later, John Anderson. 

Here it is worth while noting that in the British Museum (Natural History) a cast 
of rostrum Nr. 1884.5.3.1, River Ganges, is exhibited with the label: "the original 
specimen now in the Museum of the Royal College of Surgeons 2 was described and 
figured by Dr. Roxburgh it exhibits in great perfection the characters of the teeth 
in aged animals, which are quite different from these of the young". It is tempting 
to conclude that this is the specimen previously shown to Lebeck. 

It is not certain that the "Neue Schriften der Gesellschaft Naturforschender 
Freunde zu Berlin" was also available in Calcutta in 1801. Certainly no copy 
of this journal is included in the Indian Museum library to-day. "Asiatick 
Researches", on the other hand, must have been easily accessible to Lebeck who 
was living in India, or had lived in India, at about that time. 

There is no doubt that Roxburgh made very thorough observations of the Hughly 
river dolphins, since his house, as Anderson (1878) explained, resembled the bridge 
of a ship and commanded an extensive view of the Hughly river. Gangetic dolphins 
still swim in this river to-day as they did in the past. Roxburgh was also interested 
in hydrobiological problems (water insects) which is further proof of his familiarity 
with Indian rivers. He was considered an expert responsible for many botanical 
discoveries and descriptions of Indian plants. 

Although he was predominantly a botanist Roxburgh did have the preliminary 
training as a surgeon so, like Anderson, he would have had the basic knowledge of 
anatomy that would have made him competent to deal with a dolphin. It is incon- 
ceivable to me that a man of his experience would need to copy a report of Lebeck. 
The contrary is much more plausible. Therefore, I am of the opinion that William 
Roxburgh, and not H. J. Lebeck, was the first author to describe this interesting 
species of cetacean, and suggest that in the future publications on the Gangetic 
dolphin, the species should be referred to as Platanista gangetica (Roxburgh 1801). 



ACKNOWLEDGEMENTS 

The biographical data on William Roxburgh were placed at my disposal by my 
friend and colleague, Dr. Francis C. Fraser, British Museum (Nat. Hist.), London. 
Grateful acknowledgements is also made of the kind co-operation of the Trustees and 

1 "the second rostrum I was shown of such an animal". 

2 The specimen was destroyed by enemy action during the second world war. 



348 G. PILLERI 

the Keeper of Zoology Dr. J. P. Harding of the British Museum (Nat. Hist.), London, 
who allowed me admittance to the osteological collections. I also wish to extend my 
thanks to Dr. M. Gihr, Dr. C. Kraus, Prof. B. Horning in Berne, and Dr. U. M0hl, 
Zoological Museum Copenhagen, for their help in the bibliographical research and to 
Mrs. Diane M. von Nordheim, Geneva, for the English translation. The work was 
sponsored by the Swiss National Fund for the Promotion of Scientific Researches. 



SUMMARY 

The attribution of the original description of Platanista gangetica to Lebeck is 
contested. It is first established that the descriptions of Lebeck and Roxburgh 
were published in the same year, i.e. in 1 80 1, and the reasons for concluding that the 
description of Lebeck was copied from that of Roxburgh are set out. It is suggested 
that Roxburgh be considered the author of the original description of the Gangetic 
dolphin. 



REFERENCES 

ANDERSON, J. 1878. Anatomical and Zoological Researches: comprising an Account of the 

Zoological Results of the two Expeditions to Western Yunnan in 1868 and 1875; and a 

Monograph of the two Cetacean Genera Platanista and Orcella. 2 vols. London (B 

Quaritch) . 
CUVIER, M. F. 1836. De 1'histoire naturelle des Cetaces ou recueil et examen des faits dont se 

compose 1'histoire naturelle de ces animaux. 416 pp. Paris (De Rovet). 
CUVIER, G. 1823. Recherches sur les Ossements Fossiles de Quadrupedes, ou Ton retablit 

les characteres de plusieurs especes d'Animaux que les revolutions du Globe paraissent avoir 

detruites &c. 5 : 279-280. Paris (Dufour et D'Acagne). 

GOSCH, C. C. A. 1875. Udsigt over Danmarks Zoologiske Literatur. 2. Afd., Bd. 2, Kobenhavn. 
HERSHKOVITZ, P. 1966. Catalog of Living Whales. Bull. U.S. natn. Mus. 246 : 1-259. 
HOME, E. 1818. A description of the teeth of the Delphinus gangeticus. Phil. Trans. R. 

Soc. 108 : 417-418. 
LEBECK, H. J. 1801. Delphinus gangeticus beschrieben vom Herrn Heinrich Julius Lebeck zu 

Trankenbar. Neue Schr. Ges. naturf. Fr. Berl. 3 : 280-282. 

LEE, S. 1897. Dictionary of National Biography. 44 : 368. London (Smith, Elder & Co.). 
ROXBURGH, W. 1801. An Account of a new Species of Delphinus, an Inhabitant of the 

Ganges. Asiat. Reschs. 7 : 170-174. Calcutta. 
1803. (printed verbatim from the Calcutta edition). London. 



Professor G. PILLERI 
BRAIN ANATOMY INSTITUTE 
UNIVERSITY OF BERNE 
3072 OSTERMUNDIGEN 
SWITZERLAND 



A NEW GENUS AND SPECIES OF SUDAN 

LEECH FORMERLY CONFUSED WITH 
LIMNATIS NILOTIC A (HIRUDINIDAE S.L.: 

HIRUDINEA) 

By L. R. RICHARDSON 

SYNOPSIS 

A new genus is based on a leech from Zalingei Swamp, Sudan. It has 3 pairs of narrow 
broken lines on the dorsum, somitally repetitive supramarginal maculations, and a marginal 
light stripe; but there are 16 complete 5 -annulate somites, the posterior sucker is of moderate 
size, ejaculatory bulbs are present, and there are linear somital sense organs on the dorsal aspect 
of the posterior somites. A closely similar leech is recorded also from the Nile, near Fashoda. 

INTRODUCTION 

LEECHES with salivary gland papillae on the jaws have been found in all regions, 
essentially between 35 North and South. Of these, only leeches in the Mexican 
genus Limnobdella Blanchard 1893, the Australian genus Quantenobdella Richardson 
1969, and the present leech have 16 complete 5-annulate somites, all others (Soos, 
1969) have 15 such somites, including the leeches in the Ethiopian genus Limnatis 
Moquin-Tandon 1826 based on L. nilotica (Savigny 1822). The genus Limnobdella 
(v. Richardson, 1969 : 106, 'Potamobdella'} has a macrobdelloid pharynx with some 
muscular ridges ending independently between the jaws on the entrance to the 
pharynx, multiple small testes in each somite, the epididymis formed on both limbs 
of a simple primary loop on the anterior region of the male paired ducts, no ejacula- 
tory bulbs, the median regions bimyomeric and the female median region with an 
acaecate vagina and vaginal duct. The genus Quantenobdella described as lacking 
salivary gland papillae and having an hirudoid pharynx, is now found to have 
minute papillae and the pharynx smooth internally. Quantenobdella has simple 
saccular testes; the anterior region of the male paired duct lacks a loop and the 
epididymis is entirely posterior to the ejaculatory bulb, the two being linear in 
relationship; the median regions are bimyomeric and the female median region has 
a caecate vagina and a vaginal duct. 

In the present leech, the pharynx is hirudoid; the testes are simple, saccular; the 
anterior region of the male paired duct forms a simple primary loop with the epidi- 
dymis on the initial limb and an ejaculatory bulb on the terminal limb in a sub- 
parallel relationship; the median regions of the reproductive systems are as in 
Limnobdella. 

These differences warrant the provision of a separate and new genus for the leeches 
from the Sudan. It has been shown (Richardson, 1969) that the content of the 
former family Hirudinidae based on the genus Hirudo was unacceptably hetero- 
genous. Although separate families were provided then for Australian, Nearctic 

Bull. Br. Mus. nat. Hist. (Zool.) 21, 8 



350 L. R. RICHARDSON 

and Neotropical genera, the provision of a family for Oriental and Ethiopian leeches 
having an acaecate vagina, as in the present leech, must be postponed until the aquatic 
jawed sanguivores of these regions are more adequately known. For the time being, 
the new genus can be referred to the Hirudinidae s.l. 

Moore (1939) identified leeches in single collections from the American Museum of 
Natural History, the United States National Museum, and the British Museum 
(Natural History), all as being Limnatis nilotica. He briefly referred to them as 
resembling this species in having the annulation in no way differing from published 
accounts and with small papillae, large caudal suckers, the morphology of the crop 
caeca and of the reproductive systems as in Moquin-Tandon (1846, pi. vi, Haemopis 
sanguisuga) and Dequal (1912). The characteristics as taken from these authors are : 
a small anterior pair and a larger posterior pair of caeca on each compartment of the 
crop ; the epididymis formed on both limbs of a simple primary loop on the anterior 
region of the male paired duct which lacks ejaculatory bulbs; the two ejaculatory 
ducts enter independently into the atrium (Moore, 1939, Fig. 58) and the vagina 
acaecate with a vaginal duct of the same length as the vagina. The vaginal duct is 
shown as short by Dequal (1912, Fig. 13) and the vagina possibly caecate ; but there 
is definitely no duct according to Moquin-Tandon (1846, pi. vi, Figs 15, 17). Moore 
recorded from the specimens before him, the presence of only 45 to 60 teeth, a 
pattern of 3 pairs of continuous or broken dark dorsal 'lines', yellow marginal stripes 
and the venter immaculate as in 'typical North African' examples of this species. 

Moore's material in the British Museum (Natural History) was collected in 1925 
and came from Zalingei Swamp, former British Sudan. Specimens from this collec- 
tion and another from the Nile near Fashoda, 1907, were loaned to Keegan et al. 
(1969) who figure without other description a leech from one of these collections, 
under the name of Limnatis nilotica. (The external features, Figs 7A&B, 8A&B; 
the jaw, Figs 8 C & D, and reproductive system, Fig. 8 E.) 

The reproductive system as shown in Keegan et al. differs from the illustration in 
Moore (1939, Fig. 58) of the system in a specimen in the U.S. National Museum 
collection No. 5501, from between Abyssinia and British East Africa, in that Keegan 
et al. show small ejaculatory bulbs embracing the epididymi and the two ejaculatory 
ducts joining to form a distinct common duct leading to the atrium. No such 
common duct has been known in jawed sanguivorous or macrophagous leeches, the 
two ducts always entering the atrium independently. 

Re-examination of specimens from the 1925 Zalingei Swamp and 1907 Fashoda 
collections confirm the basic pattern as described by Moore and shown in Keegan 
et al. The jaws carry minute teeth and papillae. Three dissected leeches show 
anterior and posterior paired caeca on the crop compartments but differ from Keegan 
et al. in having the ejaculatory ducts entering independently into the atrium, as in 
Moore (1939). They further agree with Moore in having a single primary loop on 
the anterior region of the male paired duct but differ by having the epididymis on 
the initial limb of the loop subparallel to the bulb on the terminal limb, as in Keegan 
et al. They differ again from Moore in having linear somital sense organs on the 
dorsum of the posterior part of the body, as in most species of Limnatis (i.e. excepting 
paluda and possibly nilotica). 



A NEW GENUS AND SPECIES OF SUDAN LEECH 351 

In having 16 complete 5-annulate somites, the specimens differ from the genus 
Limnatis as defined by Moore (1927) and Soos (1969). As I have shown previously, 
if we attempt to follow Moore (1927 et seq.) and admit such a difference into a single 
genus, the generic entity fails on other grounds (Richardson, 1969, 1970). There is 
no established genus suitable for them. Accordingly I provide a new genus as 
below. 

DESCRIPTION 

Aliolimnatis gen. nov. 

Hirudinidae s.l. with a monostichodont condition ; 16 complete 5-annulate somites 
(ix to xxiv) ; xxv, 4-annulate ; somital sense organs, large and obvious on the dorsum, 
circular anteriorly, mostly linear and oblique posteriorly; jaws, moderate in size; 
teeth, minute, about 80; salivary gland papillae on the jaws; dorsal salivary glands, 
compact, a single mass without obvious columns of aggregated ducts; radial muscles, 
a distinct extrinsic system in viii and ix ; pharynx and associated structures, hirudoid ; 
mouth and lumen of pharynx, narrow, the lumen tubular; pharynx with six internal 
muscular ridges as dorsomedian and ventrolateral pairs, each pair joining to enter a 
jaw, none ending independently between the jaws; pharynx terminating posteriorly 
in ix; crop compartments in x to xviii each with an anterior small and posterior 
larger pairs of caeca, the posterior pair in xix forming the postcaeca extending to 
xxvi; intestine, simple tubular, joining terminally to the rectum; genital pores in xi 
and xii bs/be; testes, normally 10 pairs; anterior region of male paired duct folded as 
a simple primary vertical loop, the epididymis on the initial limb posterior to the 
ejaculatory bulb on the terminal limb, the relationship subparallel; median regions, 
bimyomeric, mesomorphic; penis sheath reflected anteriorly; oviducts, short, of the 
length of the ovaries; atrium, thick-walled, large; common oviduct, thick-walled, 
longer than the oviducts, continuous with the acaecate vagina; vaginal duct, short. 

Size, (?) medium. Pattern, longitudinal interrupted narrow dark bands, marginal 
contrast stripes. 

Aquatic. Sanguivorous. Ethiopian Region. 

The name Aliolimnatis is derived from alms another, and limnatis of the marshes. 
The gender is feminine. 

TYPE SPECIES: Aliolimnatis diver sa sp. nov. 



Aliolimnatis diversa sp. nov. 
Fig. i A-F 

HOLOTYPE: B.M.(N.H.) Reg. No. 1970 3 i 

SCHIZOHOLOTYPE : B.M.(N.H.) Reg. No. 1970.3.2 right ventrolateral jaw (micro- 
slide). 

PARATYPES: B.M.(N.H.) 1970.3.3 (18-0 mm long). 1970.3.4. 
LOCALITY: Zalingei Swamp, Sudan. (Coll. Admiral Lynes.) 



352 L. R. RICHARDSON 

GENERAL FORM. Holotype (Fig. i, E.) preserved in alcohol, rather strongly 
contracted with the annuli generally at least as high as long; short, heavy-bodied, 
depressed, the dorsum almost flat, the margins obtusely rounded, the venter flat, 
and the depth nearly uniform along the length of the body. 

The anterior sucker broad, the aperture transverse ; the velum proper contracted, 
thick margined and turned ventrally; the body widening gradually behind the 
sucker, at first subcircular in section, then widening more rapidly to the anterior end 
of the clitellum and from here to the post-nephridial region, the width uniform, the 
margins parallel, converging abruptly in the post-nephridial region to form the 
narrow base to the sucker. The posterior sucker is about half the maximum width 
of the body. 

Total length, 22-0 mm; depth generally 3-0 mm; width at iv/v, 2-4 mm, at vi/vii 
4-0, at ixjx 4-8; clitellum and testicular region, 6-0 mm wide; basis for sucker, 2-0 
and diameter of sucker 3-0 mm. 

COLOUR AND PATTERN. (Fig. i, A, B, C, E.) Preserved, faded; general back- 
ground colour, faintly yellowish brown on the dorsum, paler on the immaculate 
venter which is separated by a pair of dark longitudinal bands from distinct pale 
cream marginal stripes. A dark patch across the posterior quarter of the dorsum of 
the sucker which is otherwise pale above and below. 

Three pairs of weakly indicated maculated interrupted dark longitudinal lines 
form narrow bands on the dorsum, dividing it into a median stripe, a wide inner and 
two narrow outer pairs of longitudinal stripes, all of the background colour so that 
the dorsum is not brilliantly or colourfully striped, and the marginal stripes provide 
the only contrast colour other than the dark bands. The maculations which form 
the dark bands have the appearance of very large individual chromatophores. 

The ocular arch is on the edge of the background colour, with the 5th pair of eyes 
in from this edge which becomes lateral to the supramarginal line in viii and immedi- 
ately lateral to the line of marginal somital sense organs along the greater length of 
the body including xxvi, so defining above the cream marginal stripe continuous 
around the velum and back to xxvi/xxvii. 

The inner and middle pairs of dark bands extend along the medial and lateral 
borders of the paramedian fields; the inner pair commencing in viii and extending 
to xxvi a.2, define the median light stripe between these limits as filling the median 
field and including the paramedian sense organs. Between the ocular arch and viii, 
the median field and much of the interocular area are vaguely darker than the 
general colour elsewhere. The middle pair of narrow bands extend from in vii into 
xxvi (possibly also into xxvii for the dark patch on the sucker is divided as though 
into topographic equivalents of the inner and middle pair of bands). Between the 
inner and middle paired bands, the inner paired light stripe fills the greater part of 
the paramedian field, increasing in width as this field widens along the length of the 
body. The outer paired narrow dark bands are lateral in the intermediate field, 
extend from in viii back into xxv and between these limits define the middle paired 
stripes which include the intermediate sense organs and the greater part of the 
intermediate field. 



A NEW GENUS AND SPECIES OF SUDAN LEECH 353 

The narrow outer paired stripes are lateral to the outer paired bands, denned 
laterally by the marginal stripe, and include the supramarginal sense organs, the 
supramarginal field and marginal sense organs along the greater length of the body. 

In the middle half of the body, there are distinct small darker patches close to the 
lateral edge of the outer paired stripe. Each patch is restricted to an annulus. 
These occur with recognizable regularity on bi and bs, in some somites on b 2 and b$ 
and in a few somites on bi on only one side or the other. Such patches are present 
from viii to xxvi. The patches are spaced, nowhere give the appearance of a band, 
and form a distinct subsidiary pattern as in Hirudinaria and Poecilobdella. 

The dark band separating the marginal stripe from the venter, extends along the 
line of the submarginal sense organs and briefly into the fields on either side. 

ANNULATION. (Fig. i, A, B, C.) Preserved, contracted. Intersomital and 
interannular furrows, very deep, equivalent; no obvious division of annuli into 
couplets or triplets; somital limits not indicated generally; annuli liberally divided 
by fine longitudinal lines into small rectangles. Somital sense organs are circular 
and difficult to detect on the venter, but obvious in large clear patches on the dorsum 
where all are circular in the anterior region but progressing posteriorly: the inter- 
mediates are elongate, linear and oblique to the long axis of the body; the supra- 
marginals are linear along the long axis ; then the paramedians are linear and oblique ; 
and the supramarginals, linear and transverse. Linear somital sense organs occur 
back into xxvi ; all are circular in xxvii. Sensillae are obvious with a central sensilla 
surrounded by others forming a rosette, the rosettes arranged as a row across the 
annulus, each rosette situated in a rectangle. Nephropores are obvious, minute, and 
situated close to the posterior border of ai and b 2 just laterally to the line of the 
intermediate sense organs. 

The velum proper is broad with a thick incised margin. The velum includes the 
ist to 3rd pairs of eyes ; the first furrow iv/v does not reach the margin and the dorso- 
lateral lobe of the margin of the sucker is not strongly defined anteriorly; v, 2- 
annulate above, aia2 with the first eyes and first detectable paramedians > as, the 
furrow aia2/aa reaching into the supramarginal fields with aia2 briefly forming the 
lateral margin of the sucker which is completed by uniannulate v which also forms 
the ventral margin; vi is 3-annulate above, the 5th pair of eyes in a2, ai<a2>as, 
ai/a2 ending in the supramarginal field and vi is 2-annulate below with aia2 > as; 
vii, 3-annulate above and below, ai<a2<as; viii, 4-annulate with ai (=vii as) > 
a 2 > bs = be and with the first pair of nephropores on ai ; ix to xxiv are all 5- 
annulate (total 16) ; ix, bi = b 2 = a 2 < b 5 = b 6 ; x, bi = b 2 < a 2 = b s > b e , as 
also xi; due to contraction, the relative lengths of annuli are not assessable with 
confidence in xii to xvii; xviii, bi = b2 < a2 < bs > be, as also xx and xxi, so that 
b 5 may possibly be the longest annulus in typical 5-annulate somites of the middle 
nephric group; xxiv, bi = b 2 < a 2 > b 5 = be, and the last nephropores on b 2 ; 
xxv, 4-annulate above and below, bi =b 2 < a 2 < a 3 ; xxvi, 3-annulate above, 
ai < a2 > as but definitely 4-annulate below, ai being divided into bi < b 2 by a 
well-formed furrow which does not quite reach the margins of the body and is not 
to be seen from above; xxvi 3.3 is the last annulus across the venter and is much 



354 



L. R. RICHARDSON 



2.0 




xm 



FIG. i. Aliolimnatis divevsa gen. et sp. nov. Dorsal annulation and pattern A. somites i 
to x, B. midnephric region, and C. somites xxv to ^.wn and posterior sucker. D. Jaws, 
and internal muscular ridges of the pharynx opened along the midventral line (arrow 
marks mid-length of pharynx) . E. Ventral aspect showing general form and submarginal 
bands. F. Anterior region of male paired ducts, male median region, and female repro- 
ductive system (arrow marks dorsal aspect of male primary loop) . All figures from the 
type. Roman numerals indicate somites and somital ganglia. Somital ganglia 
represented at relative size. A. gl. albumin gland; at. atrium; c. od. common oviduct; 
ej. b. ejaculatory bulb; epid. epididymis; ov. ovary; p.s. penis sheath; pr. prostate; va. 
vagina; va. d. vaginal duct; v.d. vas deferens. Scales in mm, 0-5 mm unless otherwise 
indicated. 



A NEW GENUS AND SPECIES OF SUDAN LEECH 



355 



shortened; xxvii, incomplete 2-annulate with a brief accessory annulus. The anus 
is at the posterior border of xxvii. 

Dorsum of the posterior sucker with 6 concentric furrows; some linear sense 
organs are detectable. The ventral face has some 20 primary muscle bands centrally, 
dividing to give in the order of 50 at the margin. 

ALIMENTARY TRACT. (Fig. i, D.) The jaws are monostichodont, compressed, 
the dorsomedially wider than high at the median end with the base 0-6 mm wide, 
and the jaw 0-4 mm high; the dental margin is so low convex as to appear almost 
straight and is 0-9 mm long. There are about 79 teeth on the right ventrolateral 
jaw. The teeth are narrowly spaced, minute, about 0-018 mm tall at the median 
end, diminishing very gradually along the row so that teeth in the middle of the row 
are still 0-015 mm tall. The dorsomedian jaw is housed in an open groove, the 
ventrolaterals in open pits and the margins of all are so poorly denned that the pits 
and groove appear non-morphological. The salivary gland papillae are in three 
rows, small excepting the basal row which has some larger papillae. The dorsal 
salivary glands form a compact mass in vii to ix with no indications of columns of 
aggregated ducts. The extrinsic radial musculature of the pharynx is sparse but 
forms an obvious system in viii and ix. The pharynx commences at vi/vii. The 
entrance to the pharynx is narrow, barely wider than the base of the dorsomedian 
jaw, and the lumen is more simply tubular than tapering. The pharynx has a thin 
muscular wall with six internal muscular ridges arranged as dorsomedian and ventro- 
lateral pairs, each pair fusing before entering the base of the corresponding jaw, 
none ending independently on the margin of the entrance to the pharynx. The 
pharynx terminates in the posterior portion of ix followed by a short simple com- 
partment in ix as the first portion of the crop which has a short small compartment 
in x with an anterior small and posterior larger pairs of lateral caeca, as also on xi 
to xviii in which the compartments increase progressively in length and breadth, the 
anterior caeca remaining simple, small, and the posterior caeca increasing relatively 
in size, extending into the paramedian chamber and posteriorly to lie laterally to the 
anterior caeca of the following somite. In xix, the anterior caeca are small and 
restricted to the median chamber, the posterior pair form elongate postcaeca reflected 
in the paramedian chamber, reaching to xxvi and subdivided laterally into lobes. A 
short length of the crop completes the compartment in xix, connecting terminally at 
xix/xx to the simple tubular intestine which tapers to xxiv/xxv where it enters 
terminally into the short simple rectum. 



REPRODUCTIVE SYSTEM. (Fig. i, F.) The genital pores are at xi and xii 
The testes are simple saccular. There are normally n pairs situated inter- 
somitally at xiii/xiv to xxiii/xxiv. The vasa deferentia extend in the paramedian 
chambers to the level of xi\xii with the anterior region of each male paired duct 
folded vertically in a simple primary loop in xi. The small epididymis is tortuous 
on the initial posterior limb of the loop with the muscular ejaculatory bulb on the 
anterior terminal limb, and the relationship of the organs is sub-parallel. The 
ejaculatory bulbs are subfusiform. Muscular ejaculatory ducts extend medially 
into the median chamber to enter independently into the dorsal aspect of the male 



356 L. R. RICHARDSON 

atrium, not the ventral aspect as is usual, because the male median region is formed 
on an anteriorly directed primary loop reflecting at x/xi so that the atrium is at the 
posterior end of the procurrent limb continuous with the muscular penis sheath 
which has the procurrent limb dorsal to the terminal recurrent limb. 

The ovaries are elongate and tapering. The short oviducts join just behind 
ganglion xii to form an unusually large thick-walled atrium lined with a longi- 
tudinally rugose epithelium. The atrium tapers into the thick- walled common 
oviduct which is slightly tortuous and continues into the acaecate vagina. The 
vagina is sharply differentiated from the strongly muscular short thick-walled 
vaginal duct. 

The prostate is a very large thick cap covering the atrium and extending briefly 
along the procurrent limb of the penis sheath. The albumin glands are a thick 
investment of the atrium and of most of the common oviduct. 

MORPHOLOGY OF THE PARATYPES AND OTHER MATERIAL. Four specimens rang- 
ing from 16-0 to 25-0 mm in length. The general form is as in the holotype. The 
diameter of the posterior sucker is slightly wider than half of the maximum body 
width, av. 56 % (range, 50 % to 65 %) and most doubtfully of the width of the body 
excepting in extreme extension of the animal. Colour, as the holotype. Pattern, 
as the holotype in three specimens with the paired dark bands more pronounced, 
and all are distinctly maculate, nowhere continuous. The fourth specimen (25 -o mm) 
has only scattered sparse spaced maculations which do not conform to the topo- 
graphy of the paired bands in the others. Dorsal somital sense organs are distinctly 
linear on the posterior region of the first three specimens, but very vaguely indicated 
on the fourth (25-0 mm). Annulation, as in the holotype, excepting in the fourth 
specimen (25 -o mm) in which ii\iii and iii/iv cross the paramedian and median fields. 
In all, there are 16 complete 5-annulate somites, and xxvi is 3-annulate above and 
4-annulate below. 

The differences indicate a possible second species represented by the 25-0 mm 
specimen. 

Two SPECIMENS FROM THE NILE, NEAR FASHODA. Material from the second 
collection available to me from the British Museum (Natural History) included two 
specimens. Data on the label are: Limnatis nilotica Savigny. 1907.11.12 1/3. 
Loc : Nile near Fashoda. Pres : Loat Collection. 

The specimens are 39-0 and 49-0 mm long. Preserved, faded, general colour 
much as in A. diversa but slightly darker. Essential pattern as in A. diversa, 
excepting the paramedian field is moderately filled with many dark brownish bold 
maculations which are erratically placed and blend into the medial and lateral paired 
bands of this field. This gives the appearance of a broad band across the paramedian 
field and accentuates the median stripe so making it a prominent feature of the 
dorsum. This is much as shown in Keegan et al. (1969, Fig. 7). 

Somital sense organs obscure, but some linear oblique organs are detectable 
posteriorly on the dorsum; sensillae, arranged as a rosette. 

Annulation, as in A . diversa, with ix to xxiv complete 5-annulate (total 16) except- 
ing that xxiv a.% is incomplete briefly in the ventral median field, xxvi is simply 



A NEW GENUS AND SPECIES OF SUDAN LEECH 357 

3-annulate above and below with the annul! crossing the. venter as thin cutaneous 
folds, and xxvii, 2-annulate without an accessory annulus. 

Alimentary tract as in A. diver sa, excepting the intestine joins subterminally to 
the rectum so that there is a distinct rectal appendix about 2\ times as long as wide. 
The jaws were not studied for the dentition. 

Reproductive system, 39-0 mm specimen. General morphology and morpho- 
logical relationships as in A. diversa, the male median region reflecting at ganglion 
x; excepting that the anterior region of the male paired duct forms a posteriorly 
directed loop from xi into xii, on which the epididymis is on the recurrent limb and 
ventral to the bulb on the procurrent limb, the relationship is subparallel. 

Annulation and pattern suggest another possible separate species in Aliolimnatis, 
a question to be decided only from the study of more and preferably better material. 

ACKNOWLEDGEMENTS 

I am greatly indebted to Mr. R. W. Sims, Annelida Section, Department of 
Zoology, British Museum (Natural History) for the privilege of having this material 
for study; to Miss E. Pope, Australian Museum, for help in many ways; to Professor 
Marvin C. Meyer, the University of Maine, for his assistance with difficult literature ; 
to the Librarian, University of New England, for help with other literature, to the 
Science and Industry Endowment Fund for the loan of microscopic equipment. 

The study was collateral to researches on Australian Hirudinea assisted by a 
grant from the Nuffield Foundation. 

REFERENCES 

BLANCH ARD, R. 1891. Courtes notices sur les Hirudin6es. i. Sur la sangsue de cheval du 

nord 1'Afrique. Bull. Soc. zool. Fr. 16 : 218-221. 
DEQUAL, L. 1912. Contributo alia conoscenza degli Irudinei italiani. Arch. zool. ital. 5 (i) : 

1-14. 
KEEGAN, Hugh L., SEIICHI TOSHIOKA & HIROSHI SUZAKI. 1969. Blood sucking Asian leeches 

of families Hirudidae and Haemadipsidae. U.S. Army Medical Command, Japan. 406^ 

Med. Lab. Spec. Kept. pp. 1-130. 
MOORE, J. P. 1927. Arhynchobdellae. pp. 97-302 (in) Harding and Moore, Hirudinea. 

Fauna of British India. London. 
- 1939. Additions to our knowledge of African leeches (Hirudinea). Proc. Acad. nat. Sci. 

Philad. 90 (1938) : 297-360. 
MOQUIN-TANDON, A. 1846. Monographic de la famille des Hirudinees. Ed. 2., pp. 1-448 -f 

Atlas. Paris. 
RICHARDSON, L. R. 1969. A contribution to the systematics of the hirudinid leeches with 

description of new families, genera and species. Acta zool. hung. 15 (1-2) : 97-149. 
1970. Bassianobdella victoriae gen. et. sp. nov. (Hirudinoidea : Richardsonianidae) . 

Mem. Natn. Mus. Viet. 31 : 41-50. 
Soos, A. 1969. Identification key to the leech (Hirudinoidea) genera of the world, with a 

catalogue of the species, v. Family Hirudinidae. Acta. zool. hung. 15 (1-2) : 151-201. 

Dr. L. R. RICHARDSON 
4 BACON STREET 
GRAFTON, N.S.W. 
AUSTRALIA 



THE TYPE-SPECIES OF THE GENERA 
PHOXINELLUS, PSEUDOPHOXINUS AND 
PARAPHOXINUS (PISCES, CYPRINIDAE) 

By ETHELWYNN TREWAVAS 

Phoxinellus Heckel, 1843 : 1040 

Ax this place Heckel listed two species, both new, P. zeregi and P. alepidotus. P. 
alepidotus was described on the same page; it had already been named on p. ion, 
as an example of species having a certain type of pharyngeal teeth. P. zeregi was 
first described on p. 1063. If we were obliged to rely on this publication it would 
be allowable to have recourse to page-priority, regard P. zeregi as a nomen nudum 
on p. 1040 and adopt P. alepidotus as type-species. 

Bleeker early (i86oa : 423-4) made a formal decision in this sense; he denned 
Phoxinellus and referred to P. alepidotus as the typical European species ("de 
typische 1 europesche soort") ; he noted that P. zeregi should be placed in a separate 
genus, for which he proposed the name Pseudophoxinus. 

In the same year Bleeker (i86ob) published a "Conspectus systematis Cypri- 
norum" in which, though citing no type-species, he defined these two genera in 
terms consonant with his designation of the respective type-species in i86oa. 

In 1863 Bleeker deliberately changed his mind and designated Phoxinellus zeregi 
as "spec, typ." of Phoxinellus, citing Pseudophoxinus Bleeker as a junior synonym. 
This appears in three publications of 1863 (ig63a; 31; i863b: 263; i863c: 209) 
and in each he proposed a new genus, Paraphoxinus, with P. alepidotus as type- 
species. 

This is of course inadmissible and Jordan (1919) in 'The genera of fishes' came to 
this conclusion without citing all the evidence. He in fact recorded the proposal of 
Pseudophoxinus as in the publication here cited as i86ob, noting that no type-species 
were named in this brief conspectus and stating that they were supplied in the 
"Atlas" of 1863. He seems to have overlooked our 'i86oa' and i86ob is wrongly 
given the date 1859 (which belongs only to the first half of the same volume). 
Perhaps Jordan was relying on Bleeker's own confession, "Blkr. ol." (Bleeker, 
formerly), in the publications of 1863. I have not been able to discover in the works 
of either Heckel or Bleeker any reason for Bleeker's change of mind. 

The present note, then, confirms Jordan's view, by giving the evidence, that the 
only acceptable conclusion is the following : 

Phoxinellus Heckel, 1843, type-species P. alepidotus Heckel designated by 

Bleeker, 1860; synonym Paraphoxinus Bleeker 1863. 

Pseudophoxinus Bleeker, 1860, type-species by monotypy and original designa- 
tion Phoxinellus zeregi Heckel = Pseudophoxinus zeregi. 

1 This can only mean 'type-species'. The other sense of 'typical of the European spp.' could only be 
assumed if other European species had been known at that time, and this was not so. 
Bull. Br. Mus. nat. Hist. (Zool.) 21, 8 



360 ETHELWYNN TREWAVAS 

Bleeker in his three relevant statements of 1863 did not give the reference to his 
earlier (1860) designations and his second thoughts have unfortunately been followed 
by most authors, among them Steindachner (1882), Pellegrin (1923), Berg (1932, 
Zoogeographica), Karaman (1924), Tortonese (1938), Steinitz (1953) and Ladiges & 
Vogt (1965). Only Giinther (1868) used Pseudophoxinus as a subgenus (of Leuciscus) 
with the single species 'Leuciscus' zeregi, but he banished Phoxinellus altogether by 
using Paraphoxinus for P. alepidotus and putting Phoxinellus in the synonymy of 
both Leuciscus and Paraphoxinusl 

The taxonomy of these little circum-Mediterranean Cyrpinidae is very unsettled, 
and authors who have recognized P. alepidotus and P. zeregi as type-species of 
distinct genera have not agreed on the limits or definition of the genera, either with 
regard to each other or in relation to Pararhodeus Berg and Rutilus (especially R, 
tricolor (Lortet); but see Banarescu, 1960 : 67 and 119). The type-species of 
Phoxinellus, P. alepidotus, belongs to a group of populations characterized by various 
degrees of reduction of the squamation, associated with life in water whose course 
runs now in the open, now underground in limestone country in Yugoslavia. Even 
the specific boundaries are in doubt and the generic value of the reduction of scales 
is questionable. The problem has been stated by some of the authors mentioned 
above. 

To avoid misunderstanding I should say that I have no plans to work towards a 
solution myself. The need to consider P. zeregi in another context prompted me 
to look into the nomenclature. 



REFERENCES 

P. 1960. Einige Fragen zur Herkunft und Verbreitung der Susswasserfischfauna 

der europaisch-mediterranen Unterregion. Arch. Hydrobiol. 57 : 16-134, Suppl. maps and 

table. 
BERG, L. S. 1932. Obersicht der Verbreitung der Susswasserfische Europas. Zoogeographica 

1 : 107-208, pi. ii. 
BLEEKER, P. i86oa. Ordo Cyprini Karpers. Verh. natuurk. Ver. Ned. Ind. 7 (deel II 

no. 5) : 1-492. 

i86ob. Conspectus systematis cyprinorum. Natuurw. Tijdschr. Ned.-Ind. 20 : 421-441. 

i863a. Atlas Ichth. 3 : 31. 

i863b. Notice sur les noms de quelques genres de la famille des cyprinoides. Ver si. 
Meded. K. Akad. wet. Amst. 15 : 261-264. 

i863c. Systema cyprinoideorum revisum. Ned. Tijdschr. Dierk. 1 : 187-218. 
GUNTHER, A. 1868. Cat. Fish. Brit. Mus. VII. 

HECKEL, J. J. 1843. Ichthyologie in Russegger's Reisen vol. i : 993-1250. 
JORDAN, D. S. 1919. The genera of fishes. Part 2 (pp. i-ix, 163-284, i-xiii). 
KARAMAN, S. 1924. Pisces Macedoniae. 90 pp. (Split). 
LADIGES, W. & VOGT, D. 1965. Die Susswasserfische Europas. 250 pp. (Hamburg and 

Berlin. Paul Parey). 
PELLEGRIN, J. 1923. Poissons de la Syrie rapportes par M. Henri Gadeau de Kerville in 

Voyage zoologique d' Henri Gadeau de Kerville en Syrie (Avril-Juin 1908) : 5-37, pis. i-v. 
STEINDACHNER, F. 1882. Beitrag zur Kenntniss der Fische Afrikas und Beschreibung einer 

neue Paraphoxinus-Kri. aus der Herzegowina. Denkschr. Akad. Wiss. Wien 45 : 1-18. 



PHOXINELLUS, PSEUDOPHOX I N US AND PARAPHOXINUS 361 

STEINITZ, H. 1953. The freshwater fishes of Palestine. Bull. Res. Counc. Israel 3 : 207-227. 
TORTONESE, E. 1938. Viaggio del Dott. Enrico Festa in Palestina e Siria (1893). Pesci. 
Boll. 1st. Mus. Zool. Univ. Torino 46 (3) no. 85 : 1-48, pis. i and ii. 




Miss ETHELWYNN TREWAVAS D.Sc. 

c/o Zoology Department 

BRITISH MUSEUM (NATURAL HISTORY) 

CROMWELL ROAD 

LONDON, S.W-7 



A LIST OF SUPPLEMENTS 
TO THE ZOOLOGICAL SERIES 

OF THE BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 



1. KAY, E. ALISON. Marine Molluscs in the Cuming Collection British Museum 
(Natural History) described by William Harper Pease. Pp. 96; 14 Plates. 
1965. (Out of Print.) 3.75. 

2. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and 
Valenciennes. Pp. 180; n Plates, 15 Text-figures. 1967. 4. 

3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure of Mineralogy 
at the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125; 29 Plates, 
77 Text-figures. 1969. 4.50. 

4. HAYNES, J. R. Cardigan Bay recent Foraminifera (Cruises of the R.V. Antur) 
1962-1964. (In press.) 



Printed in England by Staples Printers Limited at their Kettering, Northants, establishment 



INDEX TO VOLUME 21 



The page numbers of principal references and the new taxonomic names are printed in bold type 



Abarenicola . . . . 108, in, 187 

Ablepharus . . . . . .252 

Acanthaster . . . . . 117 

Acanthocaris . . . . 15 

Acanthodactylus 

234-236, 237, 249, 250, 251, 252, PI. 4 
aceratus, Chaenocephalus . 53, 54, 64 

acetabulosus, Tadarida .... 48 

acetabulosus natalensis, Tadarida . . 48 

Acropora 113, 131, 132, 133, 139, 148, 160, 

161, 177, 179, 196, 198, 199, 202 



acrotis, Rhinolophus clivosus 
aculeifer, Hypoaspis 
aeruginosa, Polispilota . 
afer, Triaenops persicus . 
affinis, Praxillella . 
afra, Eunice . 
africana, Lironeca 
africana, Tadarida 
africanus, Miniopterus inflatus 



41, 



35 

96 

34 

PI. 3 
. 188 

107, 112, 157 

. 329-338 

47 

46 



Agama 



215, 218, 222-232, 249, 
250, 251, 252, Pis 1-3 
222-232, 252 
105, 107, 116, 154-155 
. 163 
223 

34 

106, 112, 114, 137 
112, 117, 129 
106, 134 

36 

359 

- 359, 360 

- 351-357 
308, 309 

106, 112, 113, 142 

36 



Agamidae 
Aglaophamus 
Aglaurides 
agnetae, Agama 
Agrius .... 
albini, Synelmis 
albolineatum, Phascolosoma 
albopicta, Eulalia . 
alcyone, Rhinolophus 
alepidotus, Paraphoxinus 
alepidotus, Phoxinellus . 
Aliolimnatis 
Allopolybothrus 
alternata, Typosyllis 
alticolus, Rhinolophus 
alticolus, Rhinolophus simulator . . 37 
amabilis, Malurus . 313, 314, 315, 317-318, 

320, 322, 324, 325 
Amaea 

Amaeana . . . 
amboinensis, Lycastopsis 
amboinensis, Namanereis 
americanus, Lithobius 
Ammotrypane 
Ampharetidae 
Ampharetinae 
Amphinome . 
Amphinomidae 
Amphiroa . . . 
ampullifera, Harmothoe 
ampullifera, Polynoe 



195 
108, 195 

147 
117, 118, 147 

297, 299, 302, 309 

. 182 

105, 108, 115, 193-194 

194 
106, 113, 130 

105, 106, 130-133 

112, 126, 131, 151 

126 

126 



ampulliferus, Paralepidonotus 
Anaitides . . 106, in, 118, 
ananas, Thelenota . 
Ancistrosyllis 

Angel, M. V. . . 257 

angolensis, Hipposideros caffer 
Angosoma .... 
Angosomaspis 
Anguidae .... 
anomala, Pseudonereis . 
antarctica, Ogmogaster . 
antennata, Eunice . 107, 

antillensis, Eunice . 
antipoda, Pectinaria 
anurus, Epomophorus 
Aphroditidae . 104, 105, 

aphroditois, Eunice 
apodus, Ophisaurus 
Arabella . . 107, 

Arabellinae .... 
arenaceodonta, Neanthes 
arenaceodonta, Nereis 
Arenicola .... 
Arenicolidae .... 
argus, Bohadschia . 
argus, Lithobius 

Armandia . . . 108, 

armillaris, Syllis 
armillaris, Typosyllis 
Asellia ..... 
asper, Acanthodactylus boskianus 
aspergillum, Cloeosiphon 
aspersa, Lacerta 
Aspidosiphon 

assimilis, Malurus 313, 314, 315, 
321, 322, 323, 324, 
assimilis, Malurus lamberti 



106, 126-127 
"9, 135-136 

117, 120 

136, 137 

-283, 289-296 

38, 39, 4 

94 

92-94, 96, 97 

253 

107, 112, 151 

54, 55 
ii2, 114, 157 

342 

108, 116, 193 

. 29 

106, 119-129 

107, 113, 157 

253 

112, 113, 164 
164-165 

. 148 
. 148 
. I8 7 

105, 108, 187 

117, 120 

303, 309 

113, 116, 182 

. 142 

106, 112, 142 

40-41 
234, 249, 252 

112, 117, 129 

234 
112, 117, 129 

316, 318, 320, 
325, 326, 327 

319 

, 86, 96, 97 
117, 120 
117, 120 
. 179 
. 96 

342 

20 



athiasae, Hypoaspis . 81-84 

atra, Halodeima 

atra, Holothuria 

Audouinia .... 

aurata, Cetonia 

auriculata, Eunice 

australiensis, Latreillia . 

australis, Taphozous perforatus . . 33 

Autolytinae ... 139 

Autolytus 1 06, 139 



Balaenoptera . . . -53, 54- 56 

balaenopterae, Diplogonoporus 54, 56 

Balanoglossus . . .113, 117, 125 
bandaensis, Sigalion . . 106, 113, 128 



364 



INDEX 



20 
25-49 

343 

3, 9, 10, 13, 16, 18 
313, 318, 320 



barbata, Homola . 
Bats .... 
benedicti, Eunice . 
benedicti, Raninoides 
bernieri, Malurus . 
bernieri, Malurus lamberti . . .319 
Bhawania . . 106, 112, 117, 129-130 

bifurcatus, Haploscoloplos 107, in, 116, 165 
bimucronatus, Promacrus ... 96 
bioculatum, Megalomma . . . 200 

bisaccata, Hymenolepis ... 53, 60 

bispinosus, Pentodon .... 74 
blanfordi, Agama .... 229, 251 
blanfordi, Ophisops .... 240 
blanfordi, Ophisops elegans 238-241, 250, 252 
blanfordi fieldi, Agama 228-230, 250, 252, PI. 3 
blanfordii, Bunopus . . . .252 
bocagei, Myotis ..... 44 
Bohadschia . . . . . 117, 120 
boskianus asper, Acanthodactylus 234, 249, 252 
Bothriocephalus .... 54, 56-58 
Bothropolys ..... 302, 309 
brachyaskos, Conchoecia . . 281, 282 

brachycola, Syllis . . . . .143 
brachycola, Typosyllis 

106, in, 112, 113, 114, 143 

brachydactyla, Agama stellio . 231-232, 252 
brachypedes, Lucanaspis 
Branchiomma 
Brania .... 
brevicirrus, Perinereis nuntia 
brevicornis, Lithobius 
brevipes, Ophiocoma 
brevirostris, Eremias 



. 88-90, 97 
108, 113, 198-199, 200 
106, 139 
. 150 
299, 303-304, 309 

117, I2O 

237 



brevirostris, Eremias brevirostris . .237 
brevirostris brevirostris, Eremias . . 237 
brevirostris microlepis, Eremias 

236-237, 2 5 2 5 2 

buchnerorum, Lithobius castaneus . 300, 308 
Bunopus ...... 252 



caducus, Dasybranchus . 

caffer, Hipposideros 

caffer, Hipposideros caffer 

caffer angolensis, Hipposideros 

caffer caffer, Hipposideros 

caffer nanus, Hipposideros 

caffer tephrus, Hipposideros . 

cantoris schmidti, Acanthodactylus 

capensis, Eptesicus 

capensis, Gyptis 

capensis, Mediomastus . 

capensis, Ophelia . 

capensis, Oxydromus 

Capitellethus 

Capitellidae . 

Capitobranchus 

Cardioderma 

carnosus, Balanoglossus . 



108, 113, 185 

38, 39, 4 
. 38-39, 40 

38, 39, 4 
. 38-39, 40 

38,39 
38,39 
252 
42 

106, 137, 138 
108, 186 

- 183 

137 
108, 184 

105, 1 08, 184-186 
108, 113, 184-185 

35 
113, 117, 125 



105, 



232- 



Carobia .... 

castanea, Carobia . 
castanea, Genetyllis 
castanea, Phyllodoce 
castaneus, Lithobius 
castaneus buchnerorum, Lithobius 
caudata, Neanthes 
caudata, Nereis 
celerio, Hippotion . 
centaurus, Angosoma 
centralis, Hipposideros ruber . 38 
Ceramodactylus 
Cerastes .... 
cerastes, Cerastes . 
Ceratonereis . . 107, 113, 

Cerithium .... 
Cetonia .... 

Chaenocephalus 
Chaetopteridae 
Chaetopterus 
Chalcides 
Chamaeleo 
chamaeleon, Lacerta 
chamaeleon musae, Chamaeleo 
chamaeleon recticrista, Chamaeleo 

232- 

Chamaeleonidae 
channeri, Lyreidus 
chestnut-shouldered wren complex 
Chilopoda .... 
chimmonis, Notosceles . 
chinensis, Leocrates 
Chiroptera .... 
Chloeia .... 

Chloephaga .... 
chloronotus, Stichopus . 
Chrysopteron 
Cichlid fishes 

cingulata, Branchiomma 108, 

cingulata, Dasychone 
cirrata, Laonice 

Cirratulidae . . . 105, 

cirratulus, Nerine . 
cirratulus saipanensis, Nerine . 
cirrifera, Prionospio 
Cirriformia . . . 107, 

claparedii, Abarenicola . 
claparedii, Abarenicola claparedii 
claparedii, Arenicola 
claparedii, Leocrates 
claparedii, Sphaerodoridium . 
claparedii claparedii, Abarenicola 
clarus, Malurus 
clavata, Brania 
clavata, Grubea 

clavigera, Gastrolepidia . 106, 116, 
clivosus acrotis, Rhinolophus . 
Cloeosiphon .... 
Clymenella . . . 108, 

coasta, Cossura 



. 136 

. 136 

1 06, 136 

106, 136 
300, 308, 309 

300, 308 

107, 148 
107, 148 

34 

94 

, 39, 40, PI. 3 

252 

253 

253 
147-148, 153 

117, 160 
. 96 
53, 54- 64 
107, 177-179 
107, 177 
241, 249, 252 
234, 249, 252 
232 

234 

234, 249, 252 
232-234, 252 

4, 9 
313-328 
297-311 

4 

106, 113, 139 
25- 4 9 

106, 116, 131 
53, 54, 58, 61 

117, I2O 

44 

329-338 

113, 198-199 

. 198 

107, 167 

107, 179-180 

174 

J 74 
107, 170 

112, 113, 179 
. 187 

108, in, 187 

. 187 

139 

107, in 
108, in, 187 

. 320 

106, 139 

139 

117, 119-I2O 

35 

112, 117, 129 

113, 187-188 

108, 180 



INDEX 



365 



coccinea, Eunice . 107, 112, 118, 119, 158 

coeliaca, Trypanosyllis . 106, in, 112, 142 

Coleolaelaps . . . 69, 76, 78, 88, 92, 96 
collaris, Eirenis ..... 253 
collaris, Lysidice . 107, 112, 114, 118, 119, 162 
colorata, Echis ..... 253 
Coluber . 215, 244-245, 249, 253, PI. 6 

Colubridae .... 243-246, 253 
commensalis, Hololepidella . . 121, 122 
complanata, Eurythoe 

106, 112, 113, 117, 130, 131 



Conchoecia . . . 
conspicua, Chloeia . 

Contracaecum . . 
convolvuli, Agrius .. 
cor, Cardioderma .. 
corallicola, Pseudopolydora 
cornuta, Ehlersia . . . 
cornuta, Langerhansia . . 
cornuta, Syllis . . 
Coronella . . 
coronella, Eirenis coronella . 
coronella coronella, Eirenis . 
coronella fraseri, Eirenis . 

corsicus, Lithobius impressus . 
corsicus, Polybothrus impressus 
Corynosoma . . . . 
Coryphaena . . . 
Cosmonotus . . . . 
Cossura 

Cossuridae . . . 
Costa, M 
costae, Ceratonereis . 



257-283, 289-296 
. 106, 116, 131 
. . 54, 64 
... 34 
... 35 
107, in, 116, 173 
. . .141 
. 106, 113, 141 
.141 
. 243 

. . . 253 
. . . 253 
. . -253 
. . . 307 
. 307 
54. 64 

53. 54. 5^ 
. 4, 9, 13, 15 
108, 180 

105, 108, 115, 180 
67-gS 
. . 107, 147 



costarum, Spiochaetopterus costarum 

107, in, 113, 178-179 
costarum costarum, Spiochaetopterus 

107, in, 113, 178-179 

costarum monroi, Spiochaetopterus . .179 
costarum okudai, Spiochaetopterus . .179 
costarum pottsi, Spiochaetopterus . .179 
coutierei, Pomatoceropsis . . . 204 
coutierei, Spirobranchus . 108, 113, 204 

cricognatha, Nereis . . . .148 

crinoidicola, Paradyte . 106, 116, 117, 126 

crinoidicolo, Polynoe . . . .126 
cristata, Pista . . . . 197 

cryptocephala, Bhawania . . 129, 130 
cryptocephala pottsiana, Bhawania . .130 
Cryptonereis . 105, 107, 117, 118, 144-147 
crypturus, Epomophorus . . .29 

cultrifera, Perinereis 107, 113, 118, 119, 149 

cumingi, Paraspidosiphon . . 117, 125 
cyaneus, Malurus ..... 325 
Cymothoidae .... 329-338 

Cyprinidae ..... 359-361 
cystops, Rhinopoma hardwickei . . 30 



Dasybranchus 
Dasychone 
debilis, Paleanotus 
decemlineata, Eirenis 
decipiens, Conchoecia 

dentata, Ophiocoma insularia 
denti, Rhinolophus 
denticulata, Ophelia 
Diadema 

diadema, Spalerosophis 
diagramma, Simochromis 
dianchora, Neofolitispa . 
dibranchis, Aglaophamus 
dibranchis, Nephtys 
dibranchis, Pista 

digitata, Thalenessa 

Diogena 

Diphyllobothrium . 

Diplocirrus . 

Diplogonoporus 

'Discovery* expeditions, 1925- 

dispar, Capitellethus 

Dispio . . . 105, 

diversa, Aliolimnatis . 

dobsoni, Rhinolophus landeri 

Dodecaceria . 

dollfusi, Mastobranchus . 

doriae, Ceramodactylus . 

dorsipes, Notopus . 

Dorveilleinae 

Dorvillea 

Drilonereis 

dubius, Hypoaspis 

dulcis, Malurus 

Dynastaspis 

Dynastes 
dysteri, Salmacina 



108, 113, 185 

. 198 

106, 112, 130 

253 
259, 269, 279, 280, 

281, 282 

117, 120 

36 



253 

329 
141, 142, 178 

154 

154 

105, 108, 113, 114, 

116, 196-197 

1 06, 129 

34 
54. 55, 56 

105, 108, 116, 181 

54- 56 

-1936 . 51-65 

. ~ . 108, 184 

107, 113, 166-167 
. 351-35 7 

35 

1 08, 112, 179-180 

108, in, 116, 185 

252 
4. 15 

. 165 

107, 165 

107, 165 

. 84-86, 97 

313, 314, 315, 317, 

321, 322, 324, 325 

90-92, 96, 97 

92 

204 



danfordi, Lacerta danfordi 
danfordi danfordi, Lacerta 



252 

252 



Eason, E. H. 
Echis . . . 
eetweldii, Eugnathichthys 
ehlersi, Euleanira . 
ehlersi, Lygdamis . 
ehlersi, Potamilla . 
ehlersi, Prionospio 
ehlersi, Sthenelanella 
ehlersi, Tetreres 
ehrenbergi, Ophisops elegans 
239, 

ehrenbergi, Terebella 
Eirenis 

elegans, Aspidosiphon 
elegans, Lepidasthenia . 
elegans, Malurus . 3 Z 3. 
elegans, Ophiarthrum 
elegans, Ophisops . 



. 297-3H 

253 
329, 337 

. 128 
108, 192 
108, 201 

107, in, 116, 170 

106, 116, 128-129 

192 

240, 241, 250, 252 
108, 112, 198 

253 

112, 117, 129 

106, 113, 117, 123 
314, 315, 322, 323 

117, 120 
238, 240, 241 



366 



INDEX 



elegans, Ophisops elegans . . 239, 252 
elegans blanfordi, Ophisops 288-241, 250, 252 
elegans ehrenbergi, Ophisops 

239, 240, 241, 250, 252 

elegans elegans, Ophisops . . 239, 252 
elegans schlueteri, Ophisops . . 239, 241 
elioti, Phyllochaetopterus 107, in, 113, 134, 177 
elongatus, Eupolybothrus . . . 308 
elongatus, Lamprologus . . . 329, 335 
elongatus, Lithobius 299, 305-3O6, 307, 309 

elongatus, Polybothrus .... 307 
elongatus impressus, Eupolybothrus . 308 

elongatus koenigi, Polybothrus . . 307 
elongatus oraniensis, Lithobius . . 307 
elongatus oraniensis, Polybothrus . . 307 



emarginatus, Henicops 
emarginatus, Lamyctes 
emarginatus, Lithobius 
enigmatica, Lironeca 
enigmatica, Mercierella 
Epomophorus 
Eptesicus 
Eremias 



erythraeensis, Ceratonereis 

eson, Hippotion 

Eteone 

Euchloron 

Euclymene 

Euclymeninae 

Eugnathichthys 

Eulalia 

Euleanira 

Eumeces 

Eunice 

Eunicidae 

Eunicinae 

Euphrosine . 

Eupolybothrus 

Eupolymnia . 

Euprepes 

Eurato 

eurystoma, Myriochele 

Eurythoe 

Euthelepus 

Euthynnus . 

exilis, Syllis . 

exilis, Typosyllis . 

eximius, Lithobius 

Exogone 

Exogoninae . 

expansus, Ichthyoxenus 

expansus, Lironeca 



297, 300 
301 
297, 300-301, 309 

329, 335. 336, 337 
108, 117, 203 
29 
42 
236-238, 249, 250, 252 



107, 147 

34 
106, in, 113, 133 

34 

108, 188 
108, 187-189 

329, 337 
106, 112, 134 

128 

241-243, 249, 250, 252, 253, PI. 5 
105, 107, 112, 113, 114, 116, 117, 
118-119, 14. 157-162, 339-344 
104, 105, 107, 157-165, 339-344 
157-162 
106, 112, 131 
306, 308, 309 
108, 114, 195, 198 
. 241 
202 

108, 116, 189-igo 
106, 112, 113, 117, 130, 131-132 
108, 114, 195 
53, 54. 55 

143 
107, 143 

300 
106, in, 112, 140 

139-140 
329, 337 
329, 335, 337 



fasciatus, Lithobius . . . 301, 309 
fausta, Diogena ..... 34 
fieldi, Agama blanfordi 228-230, 250, 252, PI. 3 
fieldi, Agama persica . . . 228, 250 



fieldi, Pseudocerastes .... 253 

filicornis, Spio .... 107, 174 

filigera, Audouinia . . . .179 

filigera, Cirriformia . . 107, 113, 179 

Filograna ..... 108, 204 

Filograninae ...... 204 

fistulicola, Dodecaceria . 108, 112, 179-i8o 

Flabelligeridae . . 105, 108, 115, 181 

flavipunctatus guweirensis, Pristurus . 252 
floricola, Potosia ..... 96 

floridana, Eunice . . . . .160 



foliosa, Euphrosine 

forficatus, Lithobius 

fraseri, Eirenis coronella 

fristedti, Phyllodoce 

fulgida, Aglaurides 

fulgida, Oenone 

fullo, Polyphylla . 

fumigatus, Rhinolophus fumigatus 

fumigatus fumigatus, Rhinolophus 

fusca, Phoebetria . 

fusca, Sabella 



106, 131 
297. 298, 302, 303, 309 

253 
106, in, 112, 134 

163 

107, 163 
78, 96 

37 

. 37 
53, 54, 62, 63 

108, 201 



fuscatus, Pipistrellus kuhlii 
fusiformis, Owenia 



42 

108, 116, 192 



Galaxea . . . 132, 134, 143, 148 

gallapagensis, Pseudonereis . . .151 
Gamasus ...... 70 

gambianus, Epomophorus ... 29 
gangetica, Platanista . . . 345-348 
Gastrolepidia . 106, 116, 117, 119-120 

Gekkonidae .... 219-222, 252 

gemmifera, Exogone . . 106, 112, 140 
Genetyllis ..... 106, 136 

Gibbs, P. E. . . . . 99-2H 

gigantea, Glycera . . 107, 113, 116, 156 

giganteus, Spirobranchus 108, 113, 203-2O4 

gilchristi, Nerinides . . . 107, 168 
glabratus, Lithobius .... 304 

Glandicephalus .... 54, 55, 56 

glandigerus, Vermiliopsis . 108, 112, 204 

Glauconycteris .... 42-43 

glaucus, Diplocirrus . . . .181 

glaucus orientalis, Diplocirrus 

105, 108, 116, 181 

Glycera . . 107, 113, 116, 156-157, 205 

Glyceridae . . 105, 107, 156-157, 205 

Glycerinae ..... 156-157 

goodei, Bhawania . 1 06, 112, 117, 129, 130 

gracilipes, Lithobius . . . .301 

gracilis, Genetyllis . ... 106, 136 

gracilis, Phyllodoce . . . 106, 136 

gracilis, Syllis . . . . 1 06, 142 

graft ei, Bohadschia . . . 117, 120 

grandiceps, Stenodactylus . . 216, 252 
grandis, Acanthodactylus . 235-236, 250, 

252, PI. 4 
grayi, Cosmonotus ... . 4 

Great Abbai Expedition, 1968 . 25-49 



INDEX 



367 



gregoryi, Notopygos . 106, in, 119, 132 

griseus, Varanus griseus .... 253 

griseus griseus, Varanus .... 253 

grossipes, Lithobius . . . .301 

Grubea ...... 139 

grubei, Eunice . . . 107, 114, 158 
guadalcanalis, Lepidasthenia 

105, 106, 113, 117, 123-125 

guineensis, Hipposideros ruber . 38, 40 

guttata, Ptyodactylus hasselquistii . .220 
guttatus, Ptyodactylus . . . .220 

guttatus, Ptyodactylus hasselquistii 

220-221, 250, 252 

guttulata, Eremias . . . -237 

guttulata, Eremias guttulata 237-238, 249, 252 
guttulata, Lacerta . . . .237 

guttulata guttulata, Eremias 237-238, 249, 252 
guttulata watsonana, Eremias . . 252 

guweirensis, Pristurus flavipunctatus . 252 
Gyanisa ...... 34 

Gyptis . . 105, 106, 114, 137-138 



haasi, Agama pallida 

215, 222-228, 250, 252, PI. i 
haedinus, Taphozous perforatus . 31-34 

Haemopis ...... 350 

Halichondria . 130, 139, 140, 141, 142, 143, 

144, 147, 148, 162, 177, 183 



Halimeda 
Halodeima . 
hamanni, Corynosoma 
hanaokai, Ancistrosyllis 
hanaokai, Sigambra 
Haploscoloplos 
Haplosyllis . 
hardwickei, Lithobius 
hardwickei, Rhinopoma 



163, 196 
117, 120 
54,64 
. 136 
116, 136 



106, in, 

107, in, 116, 165 

106, 141 

297-298, 299, 309 
30 

hardwickei cystops, Rhinopoma . . 30 
hardwickei macinnesi, Rhinopoma . . 30 
hardwickei sennaarinese, Rhinopoma 

30-31, PL 3 

hardyi, Acanthodactylus scutellatus . 251 

Harmothoe ..... 106, 120 

Harrison, C. J. 313-328 

hartmanae, Serpula . . 108, in, 203 
hasselquistii, Ptyodactylus . . 221, 222 
hasselquistii, Ptyodactylus hasselquistii . 250 
hasselquistii guttata, Ptyodactylus . .220 
hasselquistii guttatus, Ptyodactylus 

220-221, 250, 252 

hasselquistii hasselquistii, Ptyodactylus . 250 
hasselquistii puiseuxi, Ptyodactylus 

222, 250, 252 

Hemidactylus . . 219-220, 249, 252 

hendersoni, Raninoides . . . . 4, 18 
Henicops . . . . 297, 300, 301 
herdmani, Phyllochaetopterus 

107, in, 113, 117, 125, 178 
heruensis, Myriochele 105, 108, 116, 190-I9I 



Hesione 

Hesionidae . 

heterochela, Sthenelais . 

heteroclitus, Tetrabothrius 

Heteroligus . 

Hill, J. E. . 

Himerometra 

Hipposideros 

hipposideros minimus, Rhinolophus 

Hippotion 

hirsuta, Sphaerosyllis 

Hirudinaria . 

Hirudinea 

Hirudinidae, S. L. . 

Hirudo 

holobranchiata, Nothria 

holobranchiata, Onuphis 

Hololepidella 

Holothuria 

Homola 

Homolidae 



106, 114, 139 
105, 106, 137-139 

1 06, 1 1 6, 128 

54, 62-63 

88 

25-49 
117, 126 
33, 38-40, PI. 3 
35-36 

34 
106, 140 

353 
349-357 
349-357 

349 

107, 113, 116 
107, 113, 116, 163 

105, 106, 114, 116, 117, 120-123 
117, 120 
18-22 
18-24 



honiarae, Poecilochaetus serpens 

105, 107, 113, 175-176 

hunteri, Promacrolaelaps . . 94-96, 97 
Hyboscolex .... 108, 113, 181 
Hydroides . . . . 108, 112, 202 
Hydrurga . . . . 53, 54, 64 

Hymenolepis . . . 53, 54, 58-61 

Hypoaspis ..... 67-g8 

Hypsicomus . . . 108, 112, 113, 199 



Ichthyoxenus . . . . -329 

impar, Harmothoe . . . .120 

impatiens, Lumbrioconereis . . . 343 
implexa, Filograna . . . 108, 204 

impressus, Eupolybothrus . 306, 308, 309 
impressus, Eupolybothrus impressus . 308 

impressus, Lithobius . . 305, 306, 307 
impressus corsicus, Lithobius . . . 307 
impressus corsicus, Polybothrus . . 307 
impressus elongatus, Eupolybothrus . 308 

impressus impressus, Eupolybothrus . 308 

indica, Lagisca . . . . .127 
indica, Lycastis . . . . .147 
indica, Namalycastis . . 107, 117, 147 
indica, Sabellastarte .... 202 
indica, Scolelepis . . . . .168 
indicus, Lygdamis . . . 108, 192, 193 
indicus, Malacoceros . . 107, 113, 168 
indicus, Paralepidonotus . . 106, 127 

inflatum, Scalibregma . . . 108, 181 
inflatus africanus, Miniopterus . . 46 

insignis, Lithobius . . . 301, 309 

insignitus, Coluber .... 245 

insignitus, Malpolon monspessulanus 

245-246, 249, 253 

insolita, Platynereis . 107, in, 112, 150 

insularia dentata, Ophiocoma . . 117, 120 



3 68 INDEX 


integer, Coleolaelaps .... 76 


lamberti mastersi, Malurus 


319 


integer, Hypoaspis . 70, 71, 76-78, 96, 97 


Lamprologus 


329. 335, 336 


integer, Laelaps . . . . ' . 76 


Lamyctes .... 


3 01 . 39 


intermedium, Megalomma . . 108, 113 


lancadivae, Glycera . 107, 


113, 116, 156 


International Indian Ocean Expedition . 1-24 


lanceolata, Armandia . 108, 


113, 116, 182 


Iphione . . . 106, 113, 114, 123 


landeri dobsoni, Rhinolophus . 


. 35 


iracensis, Acanthodactylus tristrami . 236 


Langerhansia 


106, 113, 141 


iricolor, Arabella . . . 107, 113, 164 


Laonice .... 


107, 167 


Isolda . . . . . 108, 116, 193 


lashleyi, Diphyllobothrium 


54- 55 


isolepis, Agama . . . 229 


latastii, Ophiomorus 


253 


Isopoda . . 329-338 


latreilli, Lumbriconereis 


163 




latreilli, Lumbrineris . 107, 


114, 163-I&4 




Latreillia .... 


20, 21 


janickii, Bothriocephalus . . 54, 86-58 


leachi, Lithobius . 


. 298, 309 


japanensis, Eteone . 106, in, 113, 133 


Leachii, Lithobius . 


. 298 


japonica, Leanira . . . . .129 


Leanira .... 


129 


japonica, Magelona . . 107, 113, 175 


Leocrates .... 


106, 113, 139 


japonica, Paromola .... 20 


Lepidasthenia 105, 106, in, 113, 


117, 123-126 


japonica, Sthenolepis . . 106, 116, 129 


Lepidonotus . . . 106, 


112, 113, 126 


johnsoni, Poecilochaetus . . .176 


leptocirrus, Armandia 


108, 116, 182 


jugularis, Coluber jugularis . . . 253 


leptocirrus, Ophelina 


. 182 


jugularis jugularis, Coluber . . . 253 


Leptonychotes . . 53, 


54- 55. 56, 64 


jukesi, Lepidonotus . 1 06, 112, 113, 126 


leptonyx, Hydrurga 


53. 54. 64 


jukesi, Thormora . . 106, 112, 113, 126 


Leptotyphlopidae 


253 




Leptotyphlops 


253 




Leuciscus .... 


360 


kerguelensis, Neanthes . . . 107, 148 


leucoptera, Chloephaga picta . 


53. 54. 58, 61 


kerguelensis, Nereis . . . 107, 148 


leucopterus, Malurus 


327 


kinsemboensis, Euthelepus . 108, 114, 195 


Limnatis .... 


349-357 


kitaibelii, Ablepharus . . . .252 


Limnobdella .... 


349 


koehleri, Macrophiothrix . . 117, 123 


Limnothrissa 


329, 337 


koenigi, Lithobius ..... 305 


linaresi, Megalomma . 108, 


in, 113, 199 


koenigi, Polybothrus .... 305 


Lincoln, R. J. 


. 329-338 


koenigi, Polybothrus elongatus . . 307 


Lironeca .... 


329-338 


kollikeri, Praxilla 188 


Lithobiomorpha 


297-311 


kollikeri, Praxillella . . . .188 


Lithobius .... 


. 297-3H 


koloana, Ophelia . 105, 108, 116, 182-i83 


Lithozoea .... 


15. 23 


krameri, Gamasus ..... 70 


lobatus syriacus, Ptyodactylus 


222 


krameri, Hypoaspis . . 70, 81, 84, 96 


Loimia . . . 108, 


113, 116, 196 


kuhlii fuscatus, Pipistrellus ... 42 


Longbottom, M. R. . ". 


339-3 4 4 




longicaudata, Armandia 


108, 116, 182 




longicirrus, Scalisetosus . 


. 126 


labiata, Nycteris thebaica . . 34-35 


longipinnis, Glycera 


107, 156 


labiatus, Epomophorus .... 29 


longiseta, Hyboscolex 


108, 113, 181 


Lacerta . .219, 232, 234, 237, 241, 252 


longissima, Syllis . . 106, 


III, 112, 141 


Lacertidae .... 234-241, 252 


Lucanaspis .... 


88-90, 96, 97 


Lacertilia ..... 219-243 


Lucanus .... 


70, 96 


Lacydoniidae . 105, 107, 115, 155-156 


lucasi, Lithobius 


308, 309 


ladacensis, Coluber rhodorhachis . . 244 


lucida, Pionosyllis . 


143 


ladacensis, Zamenis .... 244 


lucida, Typosyllis . 


107, in, 143 


laddi, Dodecaceria . . . 108, 180 


Lumbriconereis 


343 


Laelaps ...... 76 


Lumbrinereis 


343 


laevigata, Naineris . . 107, 113, 166 


Lumbrinerinae 


163-164 


Lagisca ...... 127 


Lumbrineris . 107, in, 114, 116, 


119, 163-164 


lamberti, Malurus . . 313, 314, 315, 316, 


Lycastis .... 


M7 


3i8, 319, 322, 324 


Lycastopsis .... 


- M7 


lamberti, Malurus lamberti . . -319 


Lycoris .... 


. 151 


lamberti assimilis, Malurus . . -319 


Lygdamis .... 


108, 192-193 


lamberti bernieri, Malurus . . -319 


Lyreidus . . 3, 4, 7, 


9, 13, 15. 23 


lamberti lamberti, Malurus . . .319 


lyrochaeta, Aglaophamus 


155 



INDEX 



369 



lyrochaeta, Nephtys . . . 155 

Lysaretinae . . . . . .163 

Lysidice . . 107, 112, 114, 118, 119, 162 
Lysilla .... 108, 113, 116, 195 



Mabuia ..... 

Mabuya .... 243, 
macgregori, Capitobranchus 
macinnesi, Rhinopoma hardwickei . 
macintosh!, Marphysa . . 107, 
Macroclymene . . . 108, 
Macrophiothrix .... 
macrophthalma, Nudaurelea . 
Macrophyllum .... 
macroprocera, Conchoecia 



259, 269, 
278, 280, 
106, 113, 
106, 118, 
106, 114, 118, 
1 06, 



maculata, Lepidasthenia 
madeirensis, Anaitides 
madeirensis, Phyllodoce 
magalhaensis, Eulalia 
magalhaensis, Pterocirrus . 106, 
Magelona .... 107, 
Magelonidae . . . . 105, 
magna, Dispio .... 
magna group .... 

maja, Guyanisa .... 
major, Drilonereis .... 
majusculus, Triaenops persicus 
Malacoceros .... 107, 
malaitae, Gryptonereis 

105, 107, 117, 118, 

malayensis, Prionospio steenstrupi 107, 
Maldanidae . . . 105, 108, 

malmgreni, Phyllodoce . . 106, 
malmgreni, Prionospio . . 107, 
Malpolon . . 215, 245-246, 

Malurus ..... 
manihine, Eunice 
marauensis, Solomononereis 

105, 107, 151, 

maraunibinae, Gyptis 105, 106, 114, 
maritimus, Taphozous 
Markowski, S. 

maroroi, Dispio . 105, 107, 113, 

marovoi, Eunice . 105, 107, 116, 117, 
Marphysa .... 107, 
martiensseni, Otomops martiensseni 
martiensseni martiensseni, Otomops 
masalacensis, Lycoris 
masalacensis, Nereis 
masalacensis, Perinereis . 
masalacensis, Pseudonereis 107, in, 
mastersi, Malurus . . . 313, 
mastersi, Malurus lamberti 
Mastobranchus . 108, in, 113, 116, 
maxillosus, Polyodontes . 106, 

Mediomastus .... 

medusa, Loimia . . 108, 113, 

megabranchia, Eunice 



243 
249, 253 

. 185 
30 

in, 162 
"3, 187 
117. 123 

34 

134 
270-275, 
281, 282 
117, 125 
119, 135 
119, 135 
112, 134 

112, 134 
"3. 175 
197. 175 

167 
289, 291 

34 

107, 165 
41 

113, 168 

144-147 
116, 171 
187-189 
113, 135 
170-171 
249, 253 
313-328 
339-344 

152-153 
137-138 

3 1 

51-65 
166-167 

158-i6o 

in, 162 

. 46 

46 



. 119 

112, 151 
318, 320 

319 
185-186 

113, 127 
108, 186 
1 1 6, 196 

343 



megaera, Euchloron .... 34 

Megalomma . . 108, in, 113, 116, 199-201 
melanocephalus, Rhynchocalamus . . 253 
melanonotus, Polyodontes . . 106, 127 
melanops, Lithobius . . . 304, 309 
melanostigma, Sabella . . 108, 201-2O2 
meles, Heteroligus .... 88 

Melinninae ...... 193 

mendanai, Scolelepis squamata 

105, 107, 113, 174 

Mercierella .... 108, 117, 203 
Mesochaetopterus . 107, 113, 117, 123, 177 

Mesostigmata .... 67-98 

michaelseni, Lepidasthenia . . -125 
microlepis, Eremias brevirostris 

236-237, 250, 252 
microlepis, Lepidasthenia 
Micronephthys 
Micronereides 
Micronereis . 
microphyllum, Rhinopoma 
microprocera, Conchoecia 



106, 113, 117, 125 
107, 155 
146 
146 

30 

259, 264-27O, 272, 
274, 278, 280, 282 
30 



Micropteropus .... 

Millepora . . . . . .178 

minax, Hydroides . . . 108, 112, 202 
minax, Serpula ..... 202 

minimus, Rhinolophus hipposideros 35-36 

Miniopterus ...... 46 

minor, Myriochele ..... 192 

minuta, Hololepidella . . . 120, 121 
minuta, Polynoe . . . .121 

minuta oculata, Polynoe . .121 

minutus, Mesochaetopterus . . .177 
miodon, Limnothrissa .... 329 

mirabilis, Ceratonereis . . 107, 113, 148 
mirasetis, Aglaophamus . . 155 

mirasetis, Nephtys 155 

misakiensis, Notopus . 15 

moilensis, Coluber . 245 

moilensis, Malpolon . 215, 245, 249, 253 
monilicornis, Lithobius 299, 305, 306-308, 309 
monoceros, Hydroides . 202 

monoceros, Oryctes . 69, 84, 86 
monroi, Spiochaetopterus costarum . 179 

monspessulanus insignitus, Malpolon 

245-246, 249, 253 

Morris, P. . . 25 ~49 

morrisi, Chrysopteron . 44 

morrisi, Myotis . 43-46, Pis. 1-2 

Morton, J. E. .205 

mossambica, Lepidasthenia . - 123 

multidentatus, Bothropolys . 302, 303, 309 
multidentatus, Lithobius . 302, 303, 309 

multifilis, Tharyx . .180 

multisetosus, Angosomaspis . 92-94, 97 
munamaorii, Aglaophamus 

105, 107, 116, 154-155 

munamaorii, Nephtys 105, 107, 116, 154-155 
muricata, Iphione . . 106, 113, 114, 123 



37 



INDEX 



musae, Chamaeleo chamaeleon 

musculus, Balaenoptera . 

mutabilis, Agama . 

mutans, Arabella . 

Myotis . 

Myriochele 

myrtosa, Euphrosine 



234 
53, 54, 56 

. 228 
107, 112, 164 

43-46, PlS. 1-2 

105, 108, 116, 189-192 
106, 112, 131 



Naineris . . . 107, 113, 166 

najadum, Coluber ..... 253 
Namalycastis . . . 107, 117, 147 

Namanereinae .... 144-147 
Namanereis . . . 117, 118, 146, 147 

nanus, Hipposideros caffer . 38, 39 

nanus, Pipistrellus .... 42 

nasicornis, Oryctes . . 70,71, 78, 96 

natalensis, Tadarida acetabulosus . . 48 

Natrix 243, 249, 253 

Neanthes 107, 148 

nebulosa, Eupolymnia . . 108, 114, 195 
nebulosa, Polymnia . . . 195 

neglecta, Ophelia . . . . .183 

Nematonereis . . 107, 112, 113, 162 

neocaledonica, Perinereis . . . 205 

Neofolitispa . . . . 141, 142, 178 

neokrameri, Hypoaspis 71, 72, 74, 96, 97 

Neopomatus . . . . .117, 203 

Nephele ...... 34 

Nephtyidae . . 105, 107, 115, 154-155 

Nephtys . . 105, 107, 116, 154-155 

Nereidae . . 105, 107, 144-153, 205 

Nereinae ..... 147-153 

Nereis .... 107, 148, 150, 151 

Nerine ....... 174 

Nerinides 107, 168 

Newport, G ,. 297-311 

niapu, Hipposideros ruber . . 38, 40 

PNicomache .... 108, 113, 189 
Nicomachinae . . . . .189 
nigeriae, Tadarida nigeriae . . 47, PL 3 
nigeriae nigeriae, Tadarida . . 47, PI. 3 
nigricans, Harmothoe . . . 106, 120 
nigriceps, Tarbophis . . . .253 
nigrobranchiata, Amphinome . 106, 113, 130 
nigropunctata, Hololepidella 

I06, 114, 117, 120-121, 122 

nigro-punctata, Nereis . . . .150 

nigropunctata, Perinereis 107, 112, 114, 150 

nigropunctata, Polyeunoa . . .120 

nigro-punctata, Polynoe . . .120 
nilotica, Limnatis .... 349-357 

Nipa . . . . . . .117 

nobilis, Bothropolys .... 302 

norvegica, Eunice . . 107, 160-i6i, 343 

Nothria . . . 107, 113, 116, 163 

Notocirrus . . . . . .164 

Notomastus . . . . . 108, 186 

Notophyllum . . . 106, 114, 134 

Notopus ... . . 4, 9, 13, 15 



Notopygos . . 106, in, 113, 114, 118, 132 
Notosceles . . . . 4, 9, J 3> 15 

Notothenia ..... 53, 54, 64 

novaegeorgiae, Polydorella 

105, 107, 116, 168-170 
Nudaurelea ...... 34 

nudicornis, Lithobius .... 305 

nummifer, Coluber ravergieri . . . 253 
nuntia, Perinereis . . 107, 113, 149, 150 

nuntia brevicirrus, Perinereis . . .150 
Nycteris . . . . . 33, 34 



oblonga, Conchoecia . . . 259, 264 
ocellata, Lacerta . . . . .241 

ocellatus, Chalcides . . . .241 

ocellatus, Chalcides ocellatus . 241, 249, 252 
ocellatus ocellatus, Chalcides . 241, 249, 252 
octacantha, Hymenolepis ... 60 
oculata, Polynoe minuta . . .121 

oculifera, Pseudeurythoe . . 133 

Oenone ..... 107, 163 

Ogmogaster . 54, 55 

okudai, Spiochaetopterus castarum . .179 
olivieri schmidti, Eremias . . .252 
Onuphinae . . . . . .163 

Onuphis . . . 107, 113, 116, 163 

Ophelia . . . 105, 108, 116, 182-183 

Opheliidae . . . 105, 108, 182-184 

Ophelina . . . . . .182 

Ophiarthrum . . . 117, 120, 123 

Ophidia ..... 243-246 

Ophiocoma . . . . . 117, 120 

Ophiomorus . . . . . -253 

Ophisaurus . . . . . -253 

Ophisops . . . 238-241, 250, 252 

ophiuricola, Hololepidella 

105, 106, 114, 117, 121-123 

oraniensis, Lithobius elongatus . . 307 
oraniensis, Polybothrus elongatus . . 307 
Orbiniidae . . . 105, 107, 165-166 

orientalis, Acanthodactylus tristrami . 236 
orientalis, Diplocirrus glaucus 

105, 108, 116, 181 
orientalis, Homola .... 20 

Oryctes . 69, 70, 71, 78, 81, 84, 86, 96 

osculatum, Contracaecum . . 54, 64 

osiris, Hippotion . . . . .34 

Otomops ...... 46 

ovalis, Notopus . . . . .15 

Owenia .... 108, 116, 192 

Oweniidae . . 105, 108, 115, 189-192 

Oxydromus ...... 137 



Pachycephala 
pacificum, Megalomma 
palatii, Nephthys . 
palauensis, Eunice 
Paleanotus . 



326 
201 

107, 155 

342 
106, 112, 130 



INDEX 



pallida, Agama 218, 224, 225, 226, 251, Pis. 1-2 
pallida, Agama pallida 223, 228, 250, Pis. 1-2 
pallida haasi, Agama . . 215, 222-228, 

250, 252, PI. i 

pallida pallida, Agama . 223, 228, 250, Pis. 1-2 
Palmyridae . . . 105, 106, 129-130 

Palola . . 107, 112, 113, 118-119, 161-162 
paluda, Limnatis ..... 350 

pantherina, Hesione . . . 139 

papillifera, Lumbriconereis . . .164 
papillifera, Lumbrineris . 107, in, 116, 164 

Paradyte . . . 106, 116, 117, 126 

Paralacydonia . . 107, 116, 155-156 

Paralepidonotus . . . 106, 126-127 
Paraphoxinus .... 359-36 1 

Pararhodeus ...... 360 

Paraspidosiphon . . . .117, 125 

Pareurythoe . . 106, in, 113, 114, 132 

Paromola ..... 20, 21 

parthenoda, Conchoecia . 289, 293, 294, 295 
parva, Anaitides . . . 106, in, 136 
parva, Phyllodoce . . . 106, in, 136 
patrizii, Asellia ..... 41 

paucibranchiata, Pseudeurythoe 

106, 113, 116, 133 

pavimentatus, Eumeces schneideri . 242, 250 
Pectinaria .... 108, 116, 193 
Pectinariidae . . 105, 1 08, 115, 193 

pectoralis, Pachycephala . . .326 

pelamis, Euthynnus . . -53, 54, 55 
peneus, Nephele ..... 34 
pennata, Eunice . . . . -343 
Pentodon . . . . . 74, 96 

pentodoni, Hypoaspis . . 71-74, 9 6 , 97 
perfoliatus, Glandicephalus . . 54, 55, 56 
perforatus, Taphozous . 31, 32, 33, 34, 40 

perforatus, Taphozous perforatus . 31, 32, 33 
perforatus australis, Taphozous . . 33 
perforatus haedinus, Taphozous . 31-34 

perforatus perforatus, Taphozous . 31, 32, 33 
perforatus rhodesiae, Taphozous . 32, 33 

perforatus Sudani, Taphozous . . 31, 32, 33 
perforatus swirae, Taphozous . . 32, 33 

Perinereis . . 107, 112, 113, 114, 118, 

119, 149-150, 205 
Perolepis . . . . . .126 

persica, Agama .... 229, 230 

persica fieldi, Agama . . . 228, 250 
persicus afer, Triaenops . . . 41, PI. 3 
persicus majusculus, Triaenops . . 41 
personatus, Raninoides . . . . 4, n 

peruana, Prionospio . . . .173 

phaeotaenia, Hypsicomus 108, 112, 113, 199 
phalaena, Glauconycteris variegata . . 42 
phalangium, Latreillia .... 20 

Phascolosoma . . . 112, 117, 129 
philactes, Hymenolepis .... 60 

phillipsi, Leptotyphlops . . . -253 
Phoebetria . . . -53, 54, 62, 63 
Phoxinellus 359-36 1 



Phyllochaetopterus 107, in, 112, 113, 117, 

125, 126, 129, 130, 131, 134, 

135, 137. HO, ML 142, M3, 
147, 150, 151, 157, 161, 162, 

164, 177-178, 2OO, 2OI, 202 

Phyllodoce . . 106, in, 112, 113, 114, 

118, 119, 134-136 

Phyllodocidae . . 105, 106, 133-136 

phyllognathi, Hypoaspis 70, 74-76, 86, 96, 97 
Phyllognathus ..... 76 
picea, Agama stellio . . . .252 
picta leucoptera, Chloephaga . 53, 54, 58, 61 
pictum, Ophiarthrum . . .117, 123 
pictus, Polyophthalmus . . 108, 183-184 
Pilargidae . . . 105, 106, 136-137 

pilicornis, Lithobius . 298-299, 3, 39 

Filled, G 345-348 

pinnata, Prionospio . . 107, 116, 171 
Pionosyllis . . . . . .143 

Pipistrellus . . . . . .42 

Pisces 359-361 

Pista . . 105, 108, 113, 114, 116, 196-198 
pitipanaensis, Pareurythoe 

106, in, 113, 114, 132 

planci, Acanthaster . . . .117 
planus, Lithobius . . 302-303, 309 

Platanista 345~34 8 

Platynereis . . . 107, in, 112, 150 

platypus, Lithobius . . . 304, 309 
Platysphinx . . . . . .34 

pleurostigma, Lamprologus . . 329, 335 
Poecilobdella . . 353 

Poecilochaetus . 105, 107, in, 113, 175-176 
Polispilota .... -34 

Polybothrus . . . 35. 37 

Polychaeta . . 99-2 n, 339-344 

Polycirrinae . . . 195 

Polydorella . . 105, 107, 116, 168-170 

Polymnia ...... 195 

Polynoe .... 120, 121, 126 

Polynoinae ..... 119-127 

Polyodontes . . . 106, 113, 127 

Polyodontidae . . .127 

Polyophthalmus . . . 108, 183-184 
Polyphylla . . 7 8 . 96 

Pomatoceropsis . . . 204 

Porites . . 125, 130, 132, 133, 148, 

158, 163, 179, 180, 
199, 202, 203, 204 

Potamilla 108, 201 

Potamobdella 349 

Potosia .... -96 

pottsi, Spiochaetopterus costarum . 179 

pottsiana, Bhawania . . 106, 117, 130 
pottsiana, Bhawania cryptocephala . .130 

Praxilla 188 

Praxillella . . - 108, 188 

princeps, Eumeces . . . 241, 243 

princeps, Eumeces schneideri 

241-242, 250, 252, PI. 5 



372 



INDEX 



princeps, Euprepes . . . . 241 

Prionospio . . 105, 107, in, 116, 170-173 
Pristurus ...... 252 

procera, Conchoecia 259-264, 266, 268, 269, 

272, 274, 278, 280, 282 

'procera' group .... 257-283 

prolifera, Polydorella . . . 169, 170 
prolifera, Syllis . . . . .143 

prolifera, Typosyllis . . 107, 112, 143 
Promacrolaelaps . . . 94-96, 97 

Promacrus ...... 96 

propitia, Lithobius . . . .304 

proximus, Coleolaelaps .... 88 

prudhoei, Hymenolepis . -54, 58-6 1 

pruvoti, Phyllodoce . 106, in, 112, 135 

Psammolyce . . . 106, in, 113, 128 

Psammophis . . . . -253 

Pseudeurythoe . . 106, 113, 116, 133 

Pseudocerastes . . . . -253 

Pseudonereis . . 107, in, 112, 151 

pseudoparthenoda, Conchoecia . 289-2g6 
Pseudophoxinus .... 359-36 1 

Pseudopolydora . 107, in, 116, 117, 173-174 
Pterocirrus .... 106, 112, 134 

Ptyodactylus . . 220-222, 250, 252 

puiseuxi, Ptyodactylus .... 222 

puiseuxi, Ptyodactylus hasselquistii 

222, 250, 252 

pulchella, Isolda . . . 108, 116, 193 
pulcherrimus, Malurus . 313, 314, 315-316, 

322, 323, 324 

pulchricornis, Lithobius . . . 298, 299 
pumila, Tadarida .... 46-47 

punctata, Cirriformia . . 107, 112, 179 
punctatus, Pentodon .... 96 

pusillus, Micropteropus .... 30 



quadraticeps, Phyllodoce . 106, 112, 135 

quadratum, Diphyllobothrium . 54, 56 

quadrioculatum, Branchiomma . . 200 
quadrioculatum, Megalomma . 108, in, 200 
Quantenobdella ..... 349 
queenslandia, Reteterebella 108, in, 113, 198 



Ranina . 3, 4, 5, 6, 7, 9, 13, 15, 23 

ranina, Ranina . . 3, 4, 5, 6, 7, 9, 13 

Raninidae . . . . . . l-i8 

Raninoides . 3, 4, 7-11, 13, 15, 16-18, 23 

ravergieri nummifer. Coluber . . -253 
recticrista, Chamaeleo chamaeleon 

232-234, 249, 252 

recticrista, Chamaelo vulgaris . . . 232 
regularis, Perolepis . . . .126 

remanei, Dispio . . . . .167 

remilleti, Hypoaspis . . 70, 86-88, 97 
reniformis, Potamilla . . . .201 

Reteterebella . . 108, in, 113, 198 

rhinoceros, Oryctes . . . 69, 8 1 



rhinocerotis, Coleolaelaps . . 69, 78 

rhinocerotis, Hypoaspis . 78-8 1, 84, 86, 96, 97 
Rhinolophus .... 35~37 

Rhinopoma .... 30-31, PI. 3 

rhodesiae, Taphozous perforatus . 32, 33 

rhodorhachis, Coluber . 215, 244, 249, 253 
rhodorhachis, Coluber rhodorhachis 244, PL 6 
rhodorhachis, Zamenis .... 244 

rhodorhachis ladacensis, Coluber . . 244 
rhodorhachis rhodorhachis, Coluber 244, PI. 6 
rhodorhachis subnigra, Coluber . .244 
Rhynchocalamus ..... 253 

Rice, A. L. . . . . . . 1-24 

rigida, Ancistrosyllis . . . .137 

robustipinna, Himerometra . . 117, 126 
robustus, Acanthodactylus . . . 252 
rogersi, Coluber . 244-245, 249, 253, PI. 6 
rogersi, Malurus . 313, 314, 315, 317, 318, 

320, 321, 322, 324, 325 
rogersi, Zamenis . . . . .244 

Prosea, Langerhansia . . . 106, 141 
rossi, Notothenia . . . -53, 54, 64 
rothi, Eirenis ..... 253 

rouxi, Glycera .... 107, 157 

Roxburgh, W 345~34 8 

ruber, Hipposideros . . 33, 38, 39, 40 
ruber, Hipposideros ruber . . 38, 40 

ruber centralis, Hipposideros . 38, 39, 40, PI. 3 
ruber guineensis, Hipposideros . 38, 40 

ruber niapu, Hipposideros . . 38, 40 

ruber ruber, Hipposideros . . 38, 40 

rubriceps, Lithobius . . . 301, 309 
ruderata, Agama . . . . .228 
Rutilus 360 



Sabella . . . . 108, 201-202 

Sabellariidae . . 105, 108, 192-193 

Sabellastarte . . . 108, 113, 202 

Sabellidae . . . 105, 108, 198-202 

Sagittarius, Mesochaetopterus 

107, 113, 117, 123, 177 

saipanensis, Nerine cirratulus . 174 

Salmacina ...... 204 

sancti-josephi, Eurato .... 202 

sanctijosephi, Sabellastarte . 108, 113, 202 
sanguisuga, Haemopis .... 350 

savignii, Agama . 250 

savignyi, Diadema . . .117 

Scalibregma 108,181 

Scalibregmidae . . . 105, 108, 181 
Scalisetosus . . . . . .126 

schlueteri, Ophisops elegans . . 239, 241 
schmidti, Acanthodactylus cantoris . . 252 
schmidti, Eremias olivieri . . .252 
schneideri, Eumeces . . 242, 250, PI. 5 
schneideri, Eumeces schneideri 

242-243, 249, 253, PI. 5 
schneideri, Scincus .... 242 

schneideri pavimentatus, Eumeces . 242, 250 



INDEX 



373 



schneideri princeps, Eumeces 

241-242, 250, 252, PI. 5 
schneideri schneideri, Eumeces 

242-243, 249, 253, PI. 5 
schokari, Psammophis .... 253 

schusterae, Dispio . . . . .167 

Scincidae . . . 241-243, 252-253 

Scincus . . 242, 243, 253 

Scolelepis . . 105, 107, 113, 168, 174 

Scotophilus ...... 46 

scutata, Sternaspis . . . 108, 184 

scutellatus, Acanthodactylus scutellatus 

235. 236, 251 

scutellatus hardyi, Acanthodactylus . 251 

scutellatus scutellatus, Acanthodactylus 

235, 236, 251 
Selysius ...... 44 

semicincta, Audouinia . . . .179 

semisegregata, Eunice .... 343 

senegalensis, Taphozous 33 

sennaarinese, Rhinopoma hardwickei 30-31, PI. 3 
sepsoides, Sphenops . . . 251, 253 
serpens, Poecilochaetus . . . .176 

serpens honiarae, Poecilochaetus 

105, 107, 113, 175-176 

Serpula . . . 108, in, 202, 203 

Serpulidae . . . 105, 108, 202-204 

Serpulinae ..... 202-204 

serratifrons, Raninoides . . . 4, n, 18 
serrulata, Lithozoea ... 15, 23 

sibogae, Isolda ..... 193 

sibogae, Lepidasthenia . . . .126 

sibogae, Notopygos . 106, 113, 114, 132 
siciliensis, Eunice . . . 107, 112, 113, 

118-U9, 161-162 

siciliensis, Palola . . . 107, 112, 113, 

118-119, 161-162 
Sigalion .... 106, 113, 128 

Sigalioninae ..... 128-129 

Sigambra . . . 106, in, 116, 136 

silenus, Phyllognathus .... 76 

Simochromis ..... 329 

simulator, Rhinolophus . . . 36, 37 

simulator, Rhinolophus simulator . . 37 
simulator alticolus, Rhinolophus . . 37 
simulator simulator, Rhinolophus . . 37 
sinaita, Agama . 229, 230-231, 249, 252, PI. 3 
sinaitus, Hemidactylus .... 220 

sinensis, Aglaophamus . . . 155 

sinensis, Nephyts . . . . 155 

Siphonosoma . . 113, 117, 125, 126 

Sittella 326 

sloanei, Lithobius .... 300, 309 

socialis, Phyllochaetopterus 107, 112, 126, 129, 

130, 131, 135, 137, 140, 141, 142, 143, 147, 150, 

I 5 I > I 57 J 6i, l62 , 164, 178, 200, 201, 202 

Solomononereis . . 105, 107, 151-153 

somalicus, Eptesicus somalicus . . 42 

somalicus somalicus, Eptesicus . . 42 

PSosane .... 108, 116, 194 



Spalerosophis . -253 

sphaerocephala, Lumbriconereis . .164 
sphaerocephala, Lumbrineris . 107, 119, 164 
sphaerocephalus, Notocirrus . . .164 
sphaerocirrata, Micronephthys . 107, 155 

sphaerocirrata, Nephthys . . 107, 155 
Sphaerodoridae . . . 105, 107, 144 
Sphaerodoridium .... 107, in 

Sphaerosyllis .... 106, 140 

Sphenops . .251, 253 

Spio ...... 107, 174 

Spiochaetopterus . 107, in, 113, 178-179 

Spionidae . . . 105, 107, 166-174 

Spirobranchus . . 108, 113, 203-204 

splendens, Macrophyllum . . .134 
splendens, Notophyllum . 106, 114, 134 

splendida, Hesione . . 106, 114, 139 

spongicola, Haplosyllis . . . 106, 141 
spongicola, Syllis . . . . .141 

squamata mendanai, Scolelepis 

105, 107, 113, 174 
steenstrupi, Prionospio . . . .171 

steenstrupi malayensis, Prionospio 107, 116, 171 
stellio, Agama .... 232, 249 

stellio, Agama stellio .... 252 

stellio brachydactyla, Agama . 231-232, 252 
stellio picea, Agama .... 252 

stellio stellio, Agama . . . .252 

Stenodactylus . . . .216, 252 

Sternaspidae . . 105, 108, 115, 184 

Sternaspis . . . . . 108, 184 

steudneri, Tropiocolotes .... 252 

Sthenelais . . . 106, 113, 116, 128 

Sthenelanella . . 106, 116, 128-129 

sthenodactylus, Stenodactylus sthenodac- 
tylus ...... 252 

sthenodactylus sthenodactylus, Stenodac- 
tylus ...... 252 

Sthenolepis ..... 106, 129 

Stichopus . . . . . 117, 1 20 

stigmatica, Platysphinx . . . .34 

Stolonothrissa .... 329, 337 

stroemi, Terebellides . . 108, 116, 194 
stylolepis, Lepidasthenia 106, in, 113, 117, 126 
subaenea, Glycera ..... 205 

subnigra, Coluber rhodorhachis . . 244 
Sudani, Taphozous perforatus . 31, 32, 33 

swinnyi, Rhinolophus .... 36 

swirae, Taphozous perforatus . . 32, 33 

Syllidae . . . 104, 105, 106, 139 

Syllinae ..... 141-144 

Syllis . . . 106, in, 112, 141-142, 143 
Synelmis . . . 106, 112, 114, 137 

syriacus, Ptyodactylus lobatus . .222 



Tadarida 

tanganicae, Stolonothrissa 

tanganyikae, Lironeca 

Taphozous 

Tarachodes . 



46-48, PI. 3 
329 

329, 335- 337 

31-34. 4 

34 



374 

Tarbophis 
Telepsavus . 
Tentacularia 

tentaculata, Eunice . ' 
tenuis, Prionospio . 
tephrus, Hipposideros caffer 
Terebella 
Terebellidae . 
Terebellides . 
Terebellinae . 
terrareginae, Lepidasthenia 
tessellata, Coronella 
tessellata, Natrix . 
tessellata, Natrix tessellata 
tessellata tessellata, Natrix 
tetelensis, Prionospio 
Tetrabothrius 
tetraura, Lumbrinereis 
Tetreres 



INDEX 



253 
. 178 

54- 55 

107, 112, 161 

173 
38, 39 

108, 112, 198 
105, 108, 194-198 

108, 116, 194 
195-198 
. 125 

243 

243 

243, 249, 253 
243, 249, 253 

105, 107, 116, 171-173 
54. 62 

343 
192 



Thalassia . . 

Thalenessa 

Tharyx 

thebaica, Nycteris . 



125, 128, 156, 168, 178, 187 

106, 129 

108, 180 

33. 34 



thebaica, Nycteris thebaica ... 34 
thebaica labiata, Nycteris . 34-35 

thebaica thebaica, Nycteris . . 34 

Thelenota . . . 117, 120 

Thelepinae . IQ5 

Thormora . . . 106, 112, 113, 126 

tibiana, Eunice ..... 34 2 
tread welli, Prionospio . . 173 

Trewavas, E. . 359-36i 

Triaenops . . 4 1 . P'- 3 

Trichobranchinae . . . 194 

tricolor, Chrysopteron .... 44 
tricolor, Myotis . . . 43. 44. 45 
tricolor, Rutilus . . 360 

tricolor, Selysius . . -44 

tridens, Asellia tridens . 40, 41 

tridens tridens, Asellia . 40, 41 

tridentatus, Lyreidus . 3. 4. 7. 9, 13 

trilobata, Amaea . . . 195 

trilobata, Amaeana . . . 108, 195 

trinchesii, Mastobranchus 108, in, 113, 186 
trioculatum, Megalomma 108, in, 116, 200 

tristrami, Acanthodactylus . . . 236 
tristrami, Acanthodactylus tristrami 

236, 250, 252, PI. 4 

tristrami, Zootoca ..... 236 
tristrami iracensis, Acanthodactylus . 236 

tristrami orientalis, Acanthodactylus . 236 
tristrami tristrami, Acanthodactylus 

236, 250, 252, PI. 4 



Trochochaetidae 
tropicus, Poecilochaetus 
Tropiocolotes 
Trypanedenta 
Trypanosyllis 
tubifex, Eunice 
turcica, Lacerta 



105, 107, 175-176 
107, in, 113, 176 

252 
106, 142 

106, in, 112, 142 

107, 140, 161, 343 

219 



turcicus, Hemidactylus . . 219, 220, 249 
turcicus, Hemidactylus turcicus 219-22O, 252 
turcicus turcicus, Hemidactylus 219-22O, 252 
turcius, Hemidactylus . . .219 

typha, Pista . . . 108, 113, 197, 198 

typha, Terebella . .198 

Typosyllis . . . 106, 107, in, 112, 

113, 114, 142-144 
tytius, Dynastes ..... 92 



ubianensis, Lysilla 
uncinata, Dispio 
uncinata, Hydroides 
unibranchia, Pista 
unicornis, Nematonereis 
unifasciata, Neanthes 
unifasciata, Nereis 
uniformis, Exogone 
uschakovi, Mercierella 



108, 113, 116, 195 

. 167 

108, 202 

197 
107, 112, 113, 162 

. 107, 148 

107, 148 

106, in, 140 

203 



"3. "7. 
108, 



validobranchiata, Eunice 

Varanidae 

Varanus . . . . 

variabilis, Conchoecia 

variegata, Glauconycteris variegata 

variegata, Pseudonereis . . 107, 

variegata phalaena, Glauconycteris 

variegata variegata, Glauconycteris 

variegatus, Lithobius 

variopedatus, Chaetopterus 

vastus, Siphonosoma 

vermicularis, Serpula 

Vermiliopsis . 

vertagus, Cerithium 

verugera, Exogone 

vesiculosum, Branchiomma 

vesiculosum, Megalomma 

vesuvianus, Lithobius 

viaderi, Notosceles 

vinogradovi, Pista 

Viperidae j 

viridis, Eulalia .... 

vitjazi, Conchoecia 259, 269, 275-278, 

vitrarius, Spiochaetopterus 

vitrarius, Telepsavus 

vittata, Mabuia .... 

vittata, Mabuya . . . 243, 

vittatus, Scincus . 

vosseleri, Lithobius 

vulgaris recticrista, Chamaelo . 



343 

253 

253 
259, 264 

42-43 
112, 151 

42 
42-43 

302 

107, 177 
125, 126 

1 08, 203 

112, 204 

117, 160 
106, 140 

200 

108, 113, 200-201 

33 

4 

197 

253 
106, 134 
281, 282 

. 178 

. 178 

243 
249, 253 

243 

304 

232 



walhallae, Dynastaspis . . 90-92, 97 

watsonana, Eremias guttulata . .252 

weberi, Paralacydonia . 107, 116, 155-156 

weddelli, Leptonychotes . 53, 54, 55, 64 

Werner, Y. L. . . 213-256, Pis. 1-6 

wireni, PSosane . . . 108, 116, 194 

wrens, Chestnut-shouldered . . 313-328 



INDEX 375 

Zamenis ...... 244 zeregi, Pseudophoxinus .... 359 

Zelandicus, Lithobius .... 303 zeylanica, Psammolyce . 106, in, 113, 128 

zeregi, Leuciscus ..... 360 zeylanica, Sthenelais . 106, 113, 116, 128 

zeregi, Paraphoxinus .... 360 Zootoca ...... 236 

zeregi, Phoxinellus . . . 359, 360